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intJjeCitpofi^etDgorfe 

COLLEGE  OF  PHYSICIANS 
AND   SURGEONS 


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http://www.archive.org/details/pathfindersofphyOOdemp 


— From   an    engraving'. 
WILLIAM   HARVEY,   1578-1657. 


PATHFINDERS   OF 
PHYSIOLOGY 


— BY— 

J.  H.  DEMPSTER,  A.  B.,  M.  D. 

EDITOR  OF  THE   DETROIT   MEDICAL  JOURNAL;  FORMERLY  LECTURER  IN 

PHYSIOLOGY,     DETROIT   COLLEGE    OF    MEDICINE:     MEMBER   OF   THE 

DETROIT   MEDICAL   CLUB;    WAYNE   COUNTY    MEDICAL   SOCIETY; 

MICHIGAN  STATE  AND  AMERICAN  MEDICAL  ASSOCIATIONS 


Science 

'****/  taught  them  how  the  stars  do  rise 
And  set  in  mystery,  and  devised  for  them 
Number,  the  inducer  of  philosophy. 
The  synthesis  of  letters,  and  besides 
The  artificer  of  all  things,  Memory 
That  sweet  Muse-Mother. " 

—Aeschylus 


PUBLISHED  BY  THE 
THE    DETROIT    MEDICAL   JOURNAL    COMPANY 
19  14 


Published   by 

The  Detroit  Medical  Journal  Company. 

1914 


FOREWORD 

The  following-  pages  are  the  result  of  the  writer's  indulgence  in 
biography  as  a  recreation.  The  title  Pathfinders  is  presumed  to 
describe  the  contents.  The  biographical  essay,  it  is  hoped,  will  be  the 
tribute  of  a  stone  to  the  cairn  of  those  who  have  blazed  the  trail  of 
discovery  in  a  domain  that  has  meant  so  much  to  scientific  medicine, 
for  "Destiny  reserves  for  man  repose  enough."  The  writer  of  bio- 
graphy, in  his  self-appointed  task  fills  a  role  similar  to  that  of  Old 
Mortality  in  Scott's  well-known  novel  who  visited  the  graves 
of  the  departed  and  renewed  the  moss-covered  inscriptions  on 
their  gravestones.  The  chapters  which  constitute  this  volume 
have  already  appeared  in  the  Detroit  Medical  Journal  and  are 
reprinted  here  with  slight  alteration.  There  is  no  pretense  towards 
a  complete  history  of  physiology;  far  from  it.  Hence,  while  the 
courteous  reader  will  give  the  writer  credit  for  having  read,  the  criti- 
cal reader  will  discover,  perhaps,  a  great  deal  that  he  has  either  over- 
looked or  failed  to  read.  The  subject  itself  abounds  with  interest; 
regarding  the  manner  of  its  presentation,  perhaps,  not  so  much  may 
be  said.  An  endeavor  has  been  made  to  present  as  much  of  the  hu- 
man element  as  available  data  has  permitted.  Though  the  real  life 
of  every  great  man  lies  in  the  story  of  his  achievement,  rather  than 
in  the  tale  of  how  he  passed  his  days,  yet  the  human  touches  find  re- 
sponse in  the  mind  of  man.  "I  have  remarked,"  said  Carlyle,  "that  a 
true  delineation  of  the  smallest  man  and  his  scene  of  pilgrimmage 
through  life  is  capable  of  interesting  the  greatest  man;  each  man's 
life  is  a  strange  emblem  of  every  man's  and  human  portraits  faith- 
fully drawn  are  of  all  pictures  the  welcome st  on  human  walls." 


The  reader  of  the  history  of  medicine  cannot  but  be  impressed  by 
the  cosmopolitan  nature  of  the  science.  National  lines  are  unknown, 
for  thoughtful  men  of  every  clime  have  contributed  to  its  progress. 
Its  beginnings  are  enveloped  in  the  mazes  of  ancient  superstition, 
where  here  and  there  its  fitful  light  gleamed  forth  to  be  succeeded 
by  long  centuries  of  Cimmerian  darkness.  Owing  to  veneration  for 
the  work  of  such  men  as  Galen,  to  the  sacredness  with  which  the 


lifeless  body  was  viewed  and  to  the  slow  development  of  its  ancillary 
sciences  the  progress  of  medicine  up  to  the  beginning  of  the  nine- 
teenth century  was  necessarily  slow.  Medicine,  on  the  whole,  how- 
ever, has  advanced  during  periods  of  great  intellectual  activity  and 
during  times  of  intellectual  torpor  has  remained  in  a  quiescent 
state.  The  rise  and  fall  of  systems  and  methods  would  dispose 
one  to  wonder  if  the  end  is  yet;  if  we  have  at  last  reached 
the  bedrock  of  fact  in  a  scientific  sense.  The  great  advantage 
of  truth  over  error  is  that  though  at  times  crushed  to  earth, 
it  will  rise  again.  Not  until  science  and  philosophy  had  freed  them- 
selves from  the  throes  of  ecclesiasticism,  was  any  marked  forward 
movement  possible,  for,  during  the  first  fifteen  centuries  of  the  Chris- 
tian era  the  most  preposterous  ideas  of  physiology  obtained,  being 
founded  upon  the  sacred  writings  and  superstitions  of  the  saints. 
The  growth  of  knowledge  through  observation  was  scarcely  possible 
until  the  priest  was  no  longer  physician.  With  this  great  event  is 
associated  the  name  of  Hippocrates  who  was  the  first  to  make  deduc- 
tions based  upon  experiment  and  observation.  He  lived  during  the 
Golden  Age  when  Pericles  ruled  with  mild  persuasion;  when  Phidias 
made  immortal  the  sculptured  art  of  Greece  and  Herodotus  recorded 
the  history  of  the  illustrious  people ;  when  Democritus  proclaimed 
the  atomic  theory  of  the  universe  and  Socrates  taught  that  the 
greatest  knowledge  was  to  "Know  thyself."  Experiment,  observation 
and  deduction  have  been  aptly  called  the  tripod  of  science.  Though 
much  that  Hippocrates  taught  has  been  discarded,  yet  in  the  field  of 
clinical  observation  many  of  his  teachings  prevail  today.  The  "facies 
Hippocrates"  still  designates  the  characteristic  signs  of  impending 
death.  We  have  many  accurate  descriptions  of  disease  made  from 
careful  observations,  but  perhaps  more  than  all  else  we  owe  to  him 
that  lofty  idealistic  note  which  comes  down  to  us  in  the  Hippocratic 
oath. 

It  was  not  until  men  disregarded  authority  and  made  direct  appeal 
to  nature  that  medicine  experienced  its  renaissance.  Such  was  the 
method  of  Harvey,  Beaumont  and  of  others  whose  contributions  are  of 
permanent  value.  The  sincere  student  of  nature  approaches  his  subject 
with  an  open  mind ;  his  is  the  quest  for  truth.  He  possesses  "that  en- 
thusiasm for  truth,  that  fanaticism  for  veracity,  which  is  a  greater 
possession  than  much  learning;  a  nobler  gift  than  the  power  of  in- 
creasing knowledge."  As  Sir  Michael  Foster  once  said,  "His  nature 
must  be  one  which  vibrates  in  unison  with  that  of  which  he  is  in 
search ;  the  seeker  after  truth  must  himself  be  truthful,  truthful  with 
the  truthfulness  of  nature,  which  is  far  more  imperious,  far  more  ex- 
acting than  that  which  man  sometimes  calls  truthfulness."  Such  is 
the  religio  medici. 


Nor  is  the  history  of  medicine  without  its  martyrs.  While  scien- 
tific inquiry  has  been  the  chief  instrument  in  producing  a  higher  and 
better  civilization,  it  has  met  at  almost  every  step  determined  op- 
position from  the  powers  of  ignorance  and  jealousy.  There  is  great 
satisfaction  in  giving  to  the  world  those  things  which  all  men  see  and 
for  which  all  men  are  grateful.  The  poet,  the  painter,  the  musician 
and  the  architect  vie  with  one  another  in  their  appeal  to  the  esthetic 
sense.  Yet  is  there  not  something  higher  even  than  knowledge  for 
the  sake  of  knowledge,  or  art  for  art's  sake?  Yes,  there  is  honor 
to  him  who  chooses  a  less  spectacular  calling,  to  him  who  applies 
scientific  knowledge  to  the  conquest  of  disease.  Such  men  have  bat- 
tled with  the  enemy  unencouraged  by  the  blare  of  trumpets  or  the 
throb  of  the  war  drum.  They  have  pursued  their  work  in  hospital 
ward  or  laboratory,  or  as  "Weelum  McLure,"  have  braved  the  winter 
storm  on  errands  of  mercy  to  the  suffering. 


"Speak  History!     Who  are  life's  victors?       Unroll  thy  long 

annals  and  say; 
Are  they  those  whom  the  world  calls  victors  who  won  the 

success  of  the  day? 
The  martyrs  or  Nero?       The  Spartans  who  fell  at  Ther- 
mopylae's tryst, 
Or   the  Persians   and  Xerxes?     His   Judges,   or   Socrates? 
Pilate  or  Christ?" 

J.  H.  D. 


Among  the  works  by  which  the  writer  has  been  assisted  and  to  which  his 
grateful  acknowledgments  are  due  are  the  following:  William  Harvey,  by  D'Arcy 
Power;  Biology  and  it's  Makers,  by  Locy;  Lectures  on  the  History  of  Physiology 
and  Claude  Bernard,  by  Sir  Michael  Foster;  Harvey's  Work  on  the  Circulation, 
Sydenham  Society  Edition;  Beaumont's  Work  on  Digestion  (original  copy);  Life 
and  Letters  of  William  Beaumont,  by  Myer;  Brain  and  Personality,  by  Thomp- 
son; Recent  Progress  of  Heredity,  Variation  and  Evolution,  by  Locke;  Heredity, 
by  Thompson;  Gorton's  History  of  Medicine;  The  Relation  of  Medicine  to  Philos- 
ophy, Moon. — Alabama  Student,  by  Osier. 


CONTENTS 

Frontispiece  William  Harvey,  Portrait 

CHAPTER  I. 

The  Circulation  of  the  Blood— William  Harvey 1 


The  Renaissance — Anatomy  and  Physiology,  Galen  (1) — Vesalius  (2)  — 
Harvey,  birth  and  education  (3) — Fabricius  and  Harvey,  friends;  Ana- 
tomical Teaching  previous  to  1745  (4) — Harvey's  personal  characteristics 
(5) — Harvey  as  Lecturer;  His  Lecture  Precepts  (7) — Harvey  and  Bacon; 
Publication  of  the  Work  on  the  Circulation  (8) — The  Treatise  on  the  Cir- 
culation (10) — Capillary  Circulation  (12) — Asellius  and  the  Lymphatic 
Circulation  (13) — Asellius  opposed  by  Harvey  (14) — The  Lacteals,  dem- 
onstrated by  Johannes  Pecquet   (15). 

CHAPTER  II. 

Physiology  of  Digestion  in  the  Seventeenth  and 

Eighteenth  Centuries 16 

Stahl  and  Boerhaave  attacked  the  Chemical  Problems  of  Physiology  (16) 
— Peyer  and  Brunner;  Mechanical  and  Chemical  Views  of  Digestion  (17) 
— Borelli  and  Sylvius  (18) — Haller's  Elementa  Physiologia  appeared 
1757  (19) — Reaumur  and  His  Methods;  Experiments  with  Gastric  Juice; 
Spallanzani  (21) — Work  of  Reaumur  and  Spallanzani  confirmed  by  Stevens 
of  Edinburgh   (22). 

CHAPTER  III. 

Physiology  of  Digestion — William  Beaumont 23 

The  Investigations  of  Beaumont  on  the  gastric  juice  of  St.  Martin  freely 
quoted  in  Medical  Literature;  Beaumont,  His  Early  Life  (23) — The  Rou- 
tine of  a  Medical  Apprentice  at  the  beginning  of  last  century  (24) — As- 
sistant Army  Surgeon  (25) — Beaumont's  Diary  (25) — St.  Martin's  Acci- 
dent (26) — Beaumont  conceives  the  idea  of  experimenting  on  St.  Martin 
(27) — ^Beaumont  Honored  by  the  Michigan  Medical  Society  (28) — Seeks 
Assistance  of  two  Leading  Scientists  (29) — St.  Martin  Attains  Fame 
Through  His  Stomach  (30) — Beaumont  resigns  from  the  Army  (31)  — 
His  Death   (31)— His  Work  (32). 


CHAPTER  IV. 

Glycogenic  Function  of  the  Liver — Vaso  Motor  Nerves 
— Claude  Bernard 35 

Bernard's  Early  Life  and  Education  (35) — His  Productive  Period  (36)  — 
Work  on  Gastric  Digestion  (37) — Glycogenic  Function  of  tlie  Liver  (37) 
— Vaso-Motor  Nerves  (39) — The  Action  of  Carbon  Monoxide  Gas  (39)  — 
A  Friend  of  Pasteur  (40) — Domestic  Troubles  (40) — Made  One  of  the  "Im- 
mortals" (40) — His  Dexterity  as  Experimenter  (41) — ^Death  in  1878, 
State  Funeral  (41). 

CHAPTER  V. 

Respiration   42 

Views  of  the  Ancients  on  Respiration  (42) — Mechanics  of  Respiration  (42) 
— Boyle  and  His  Work  (43) — Robert  Hooke  (43) — Mayow  and  His  Re- 
searches (44) — Respiration  Prior  to  the  Eighteenth  Century  (45) — The 
Eighteenth  Century  School  (46) — Priestly  and  His  Dephlogisticated  Air 
(47) — Priestly  and  Benjamin  Franklin  (48) — Priestly  Comes  to  America 
(48) — The  Phlogiston  Theory  (49) — Lavoisier  and  His  Work  (49) — The 
First  to  Use  the  Word  Oxygen  (49). 

CHAPTER  VI. 

The  Nervous  System  50 

The  Heart  the  Seat  of  the  Soul  thought  the  Ancient  Hebrews  (50) — The 
Alexandrian  School  (50) — Galen  Proclaimed  the  "Brain  to  be  the  Seat  of 
Thought  and  Sensation"  (51) — Thomas  Willis  (51) — Frances  Glisson  Dis- 
covers Irritability  of  Muscle  (52) — Goll  and  Phrenology  (52) — Bell  and 
Magendie  (53) — Broca  and  the  "Speech  Center"  (54) — Pathologic  States 
of  the  Brain  and  Nervous  System  (55) — Tuke,  Benjamin  Rush,  and  Pinel 
(55). 

CHAPTER  VII. 

The  CeU  Theory 56 

Anticipated  iu  the  Seventeenth  Century  (56) — Bichat  and  His  Contribu- 
tion to  the  Theory  (56) — The  Theory  Announced  in  1838  (57) — Schleiden 
and  Schwann  (57) — Johann  Muller  (5  8) — Vitalism  (59) — Virchow's  Eu- 
logy on  Muller  (60) — Years  of  Discovery  (58) — Virchow  and  "Cellular" 
Pathology  (60) — The  Discovery  of  Protoplasm  by  Dujardin  (61) — Proto- 
plasm Defined  by  Starling  (62) — The  Cell  Theory  at  the  Present  Time 
(62) — The  Nucleus  (63) — Illustration  of  the  Cell  and  Cell  Division  (64)  — 
The  Cell  in  Heredity  (65). 


''There  is  no  knowledge  so  useful  to  man  as  knowledge  of 
himself.  Health  and  happiness  are  promoted  by  it.  Before 
the  advent  of  the  modem  scientific  spirit,  biologic  knowledge 
was  required  to  conform  to  the  dominant  superstitions  of  the 
time.  The  human  body  was  regarded  as  a  peculiar  and  awful 
thing,  and  not  amendable  to  the  laws  which  govern  the  rest 
of  the  universe.  Then  it  was  found  that  the  mechanics  of  the 
body  are  entirely  reconsilable  with  the  principles  of  physics. 
Humanity's  debt  of  gratitude  is  incalculably  great  to  those 
men  who  at  the  risk  of  their  lives  and  fortunes  made  dissec- 
tions of  dead  bodies  of  men  and  animals,  and  discovered  the 
mechanism  of  the  muscular  system  which  imparts  motion  to 
the  joints,  the  valvular  and  pump-like  arrangement  of  the 
heart,  and  the  hydraulic  principles  of  the  tubes  which  convey 
the  blood  through  the  body.  Then  came  those  students  of  the 
secrets  of  nature  who  discovered  that  the  same  laws  which 
govern  man  govern  the  lower  and  the  lowest  of  creatures ;  that 
between  soil  and  mineral,  fluids  and  gases,  plants  and  animals, 
there  is  no  dividing  line;  that  the  lily  is  the  daughter  of  the 
pool,  and  the  man  is  the  brother  of  the  ox.  This  knowledge 
was  gotten  for  us,  not  by  the  philosopher  among  his  books, 
but  by  the  patient  investigator  who  went  to  the  heart  of 
nature  and  studied  her  secrets." — J.  P.  Warbasse. 


CHAPTER    I. 


THE  CIRCULATION  OF  THE  BLOOD— WILLIAM  HARVEY 

"This  man  lived  in  an  age  when  alchemy  was  more  popular  than  science, 
and  the  love  of  mystery  stronger  than  the  love  of  philosophy." — Gorton. 

"History  is  simply  the  biography  of  the  mind  of  man;  and  our  interest  in 
history,  and  its  educational  value  to  us,  is  directly  proportionate  to  the  complete- 
ness of  our  study  of  the  individuals  through  whom  this  mind  has  been  manifested. 
To  understand  clearly  our  positions  in  any  science  today,  we  must  go  back  to 
its  beginnings,  and  trace  its  gradual  development,  following  out  our  laws,  difficult 
to  interpret  and  often  obscured  in  the  brilliancy  of  achievements — laws  which 
everywhere  illustrate  this  biography,  this  human  endeavor,  working  through  the 
long  ages;  and  particularly  is  this  the  case  with  that  history  of  the  organized 
experience  of  the  race  which  we  call  science." — Sir  William  Osier. 

The  Renaissance — The  renaissance,  that  transitional  movement 
in  Europe  between  the  mediaeval  and  modern  world,  affected  medicine 
and  the  sciences  at  a  much  later  date  than  art  and  letters.  It  began 
with  Petrarch  and  the  humanists  in  the  fourteenth  century  in  Italy, 
where  it  became  manifest  in  painting  and  sculpture.  The  movement 
was  accelerated  in  the  sixteenth  century  by  the  capture  of  Constanti- 
nople by  the  Turks  in  1509,  and  the  dispersion  of  its  Greek  scholars 
to  the  shores  of  Italy,  which  event  opened  anew  the  science  and  learn- 
ing- of  the  ancient  world  at  an  hour  when  the  intellectual  energy  of 
middle  ages  had  reached  its  ebb.  It  is  significant  to  note  that  Flor- 
ence, so  long  the  abode  of  intellectual  freedom  and  art,  welcomed  with 
extended  arms  the  exiled  Greek  scholars.  Her  traders  returned  from 
the  East  with  ancient  manuscripts  as  the  most  valuable  portion  of 
their  merchandise.  But  we  are  more  immediately  concerned  with  the 
movement  as  it  affected  medicine  and  its  allied  studies.  However 
much  the  new  learning  promoted  literature  and  art,  its  influence  was 
anything  but  favorable  to  the  progress  of  science.  Admiration  for 
the  literature  of  ancient  Greece  while  it  engendered  a  love  for  poetry, 
history  and  philosophy,  had  a  similar  effect  in  promoting  a  spirit  of 
veneration  for  the  writings  of  Hippocrates,  Ptolmey  and  Galen,  so 
that  it  became  almost  an  act  of  impiety  to  question  their  teachings. 
It  was  not  until  the  sixteenth  century,  as  we  shall  see,  that  the  spell 
of  ancient  authority  was  broken  by  the  direct  appeal  to  nature.  It 
was  not  until  then  that  the  anatomist  determined  at  all  cost  to  exam- 
ine the  human  body  for  himself  and  to  be  guided  by  his  own  obser- 
vations. 

Anatomy  and  Physiology — As  anatomy  precedes  physiology,  in 
order  to  adequately  appreciate  the  work  of  Harvey,  a  brief  account 
of  the  progress  in  anatomy  is  necessary.  The  great  anatomist  of  an- 
tiquity, who  surpassed  all  others,  was  Galen  (130-200  A.  D.).  He 
lived  for  a  time  at  Pergamos  and  for  five  years  at  Rome.  He  was  a 
man  of  talent  both  as  observer  and  writer.  His  writings  embody  all 
the  important  anatomical  discoveries  of  his  predecessors,  enriched  and 
much  enlarged  by  the  results  of  his  own  originality.  His  observations, 
however,  were  made  upon  the  lower  animals  on  the  faith  of  which  he 


2  PATHFINDERS  OF  PHYSIOLOGY 

expounded  the  human  subject.  Huxley  declares  that  "No  one  can 
read  Galen's  works  without  being  impressed  with  the  marvelous  ex- 
tent and  diversity  of  his  knowledge  and  by  his  clear  grasp  of  those 
experimental  methods  by  which  alone  physiology  can  be  advanced." 
Rome  was  the  field  of  his  greatest  triumph  as  physician.  Sq  great 
was  Galen's  influence  that  for  more  than  a  thousand  years  his  works 
held  undisputed  sway  over  anatomical  teaching  until  a  greater  name 
arose  in  the  person  of  Vesalius.  Vesalius,  born  in  Brussels  the  last 
day  of  1514,  inherited  from  an  ancestry  of  learned  men  a  keen  appe- 
tite for  scientific  learning.  His  was  that  independent,  liberty-loving 
mind  which  has  characterized  his  countrymen  before  and  since  his 
day.  The  great  importance  of  his  work  lies  in  the  fact  that  he  over- 
threw adherence  to  authority  as  a  means  of  arriving  at  truth  and 
employed  instead,  observation  and  reason.  Slavish  obedience  to  author- 
ity characterized  the  thought  and  methods  of  the  Dark  Ages.  This 
was  in  accord  with  the  ecclesiastical  influence  dominant  during  this 
long  period.  It  was  the  method  of  the  theologian,  which  had,  un- 
fortunately, survived  almost  to  our  own  day.  Darwin  was  perhaps 
•  he  most  recent  object  of  theological  invective.  As  the  Scriptures 
were  an  infallable  guide  to  spiritual  truth,  so  the  works  of  Galen  were 
unfailing  guides  to  scientific  truth.  Vesalius  was  bitterly 
opposed  not  only  by  the  ecclesiastic  forces,  but  by  medical  men 
of  his  time.  The  theologians  opposed  him  because,  among  other 
things,  he  differed  from  the  widely  accepted  dogma  that  man  should 
have  one  less  rib  on  one  side  because  according  to  Scripture  Eve  was 
formed  from  one  of  Adam's  ribs.  He  was  also  at  variance  with  them 
on  the  subject  of  the  Resurrection  bone.  Vesalius  was  willing,  how- 
ever, to  leave  the  matter  with  the  theologians,  since  it  did  not  appear 
to  him  to  be  an  anatomical  question.  Sir  Michael  Foster  writes  that 
Vesalius  "Tried  to  do  what  others  had  done  before  him — he  tried  to 
believe  Galen  rather  than  his  own  eyes,  but  his  eyes  were  too  strong 
for  him ;  and  he  cast  Galen  aside  and  taught  only  what  he  could  see 
and  what  he  could  make  his  students  see,  too.  Thus  he  brought  into 
anatomy  the  new  spirit  of  the  time,  and  especially  the  young  men  of 
the  time  answered  with  a  new  voice."  It  is  said  that  students  flocked 
to  his  lectures,  his  audience  amounting  to  some  five  hundred.  The 
history  of  anatomy  precedes  that  of  physiology  as  a  logical  sequence. 
The  work  of  Vesalius  placed  the  structure  of  the  human  body  in  a 
new  light. 

William  Harvey  was  the  first  man  to  study  and  proclaim  the  func- 
tion of  structures  which  Vesalius  had  in  such  a  masterly  manner 
demonstrated. 

"The  work  of  Harvey,"  says  Locy,  "Was  complemental  to  that  of 
Vesalius  and  we  may  safely  say  that,  taken  together,  the  work  of 
these  two  men  laid  the  foundations  of  the  modern  method  of  inves- 
tigating nature.  *  *  *  in  what  sense  the  observations  of  the  two  men 
were  complimental  will  be  better  understood  when  we  remember  that 
there  are  two  aspects  in  which  hving  organisms  should  always  be 
considered  in  biological  studies;  the  first,  the  structure,  and  then 
the  use  that  the  structures  subserve." 

The  new  learning  spread  over  Europe  in  a  westerly  and  northerly 
direction.  England  was  the  last  to  partake  of  its  benign  blessing. 
England  had  but  two  universities — Oxford  and  Cambridge;    France 


WILLIAM  HARVEY  3 

had  six;  Germany  eight;  Italy  sixteen.     Medicine  was  a  prominent 
department  in  all  of  them.     Compared  with  the  reception  accorded 
literature  and  philosophy,  science  lagged  in  England.     Green  sums 
up  the  situation   (1645) :     "Bacon  had  already  called  men  with  a 
trumpet  voice  to  such  studies.    But  in  England,  at  least,  Bacon  stood 
before  his  age.    The  beginnings  of  physical  science  were  more  slow 
and  timid  there  than  in  any  country  of  Europe.    Only  two  discoveries 
of  any  real  value  came  from  English  research  before  the  Restoration 
— the  first,  Gilbert's  discovery  of  terrestial  magnetism,  in  the  close  of 
Elizabeth's  reign;  the  next,  the  great  discovery  of  the  circulation  of 
the  blood  which  was  taught  by  Harvey  in  the  reign  of  James.    Apart 
from  these  illustrious  names  England  took  little  share  in  the  scien- 
tific movement  of  the  continent;  and  her  whole  energies  seemed  to 
be  whirled  into  the  vortex  of  theology  and  politics  by  the  Civil  War." 
Birth  and  Education — William  Harvey  was  bom  in  Folkstone, 
England,  April  1st,  1578.    Very  little  is  known  of  his  early  life.    His 
preliminary  education  was  obtained  at  his  native  town,  where  he 
made     his     first     acquaintance     with     Latin.       He     proceeded     to 
the  King's  School,  Cambridge,  where  he  remained  five  years,  and 
afterward,  at  16  years  of  age,  entered  Caius  College,  Cambridge,  in 
1593.    Harvey  even  early  in  his  school  life  possessed  habits  of  minute 
observation.    His  fondness  for  dissections  and  his  love  for  compara- 
tive anatomy  had  shown  his  mental  bias  from  his  earliest  years.    To 
Caius,  the  founder  of  the  College  at  Cambridge,  is  accredited  the  in- 
troduction into  England  of  the  study  of  practical  anatomy.     He  ob- 
tained for  his  college  a  charter  which  allowed  the  authorities  of  the 
institution  to  take  annually  the  bodies  of  two  criminals  condemned  to 
death  and  executed  at  Cambridge,  free  of  all  charges,  for  the  purposes 
of  dissection,  with  the  view  to  increase  the  knowledge  of  medicine 
and  to  benefit  the  health  of  her  majesty's  lieges,  without  interfer- 
ence on  the  part  of  any  of  her  officers.    To  what  extent  the  college 
availed  itself  of  the  privilege  is  not  known.     In  all  probability  Har- 
vey pursued  the  course  of  study  which  consisted  of  a  sound  knowledge 
of  Greek  and  Latin  ordinarily  followed  until  he  obtained  his  B.  A. 
degree  in  1597.    A  year  after  graduation,  at  the  age  of  twenty,  we 
find  him  traveling  on  the  continent  where  he  studied  the  scientific 
branches  tributary  to  medicine,  as  well  as  medicine  itself.     As  has 
been  said,  the  universities  of  northern  Italy  were  the  first  to  welcome 
the  new  learning  as  it  emanated  from  the  east  in  the  minds  of  Greek 
scholars,  as  well  as  rescued  manuscripts.    The  universities  of  north- 
ern Italy,  namely,  Bologna,  Padua,  Pisa  and  Pavia,  were  at  the  time 
at  the  height  of  their  renown  as  centers  of  mathematics,  law  and 
medicine.    Harvey  studied  more  particularly  at  Padua,  renowned  for 
its  anatomical  school,  and  rendered  famous  by  the  work  of  such  men 
as  Vesalius,  the  first  of  modem  anatomists,  and  his  successor,  Fabri- 
cius.    The  tolerance  shown  towards  Protestants  in  Padua,  the  univer- 
sity town  of  Venice,  the  great  commercial  republic,  attracted  many 
law  and  medical  students  from  England  and  other  Protestant  coun- 
tries of  Europe. 

It  is  interesting  to  recall  that  each  entry  in  the  university  (Pa- 
dua) register  was  accompanied  by  a  note  describing  some  physical 
pecularity  of  the  student,  as  a  means  of  his  identification.  Thus 
Johannes  Cookaeus,  Anglus  cum  cicatrice  in  articulo  medii  digiti  die 


1»  :..  -. 

.-:    .. 

.   T- 

and 

Z  c. 

.1  I'l 

rl. 

i'=-i 

>-  - 

■'  -■ 

■  ^-- 

thr-. 

rl'r 

4  ?AT:--r::oz?.5  or  phtsiolosy 

dicta.  John  Cotdc,  an  Rngtishman,  with  a  sear  over  the  joint  of  his 
middle  finger.  (Matncniated)  on  the  same  day,  and  so  on.  Hairey 
evidently  did  not  enter  Padua  Uniirersi^  as  a  r^nlar  matriealant, 

as  no  such  Tccord  rj^z^nrs:  on  the  Tizii-rersity  r^rister  r^ardiog  him. 

Fabricins  and  Har"vey  Friends- — Thr  fame  of  some  of  its  medical 
:rs::  ri^      ::  T         ?  ::;::   f  H-:    t     :     Padna.     While  there  he 

-  £=  mi :: .  :  r  ?:::;;    ai.i  ^lys  ;  ;r       7  Fabridns,  one  of  the 

-:::  \'.-  -rai:  r       :;;.  :^   ::  I:a-7.     i::t  :a:::r  a=  ai.atamist  and  smgeon 
::   7ah:::        ?      ^      ?^r:;iT:i:e     i:   -  jhr   :  a— r    :i  his  hirthplace) 
-'^.       ri  z.   :    ;t      _.:::.  ri;:    ^    ^  ^ :   :  uro  in  Padna  he 

^roai-i^r  fa^:  fi'ir:::.?,  A:  :J:a:  ::arT::u" j,r  Time  Fahricias 
'::.  priZT;::::^-  hi;  !£::;  ^^irr  ::  :hr  ;^/ rs  :f  the  veins. 
v>         T  r     a.  t3;  31:3     ::t      :     :-i  -rz:  ::;   ::  rhe  ves- 

'\.\\'\  ^a-V  =  T::  :r:i;i  :hT  larrr  ::  :hr  =:::aLT:'  "rizs,  while 
:  rrzuirri  :::  :hr  arTri'ir^  brca/i^r  "hi  :.::a  ~as  a'~ays 
in  a  iTazr  ■::  ra:  r.a::  :_:':'a  Har"'T7.  ai^^rvar,  ^::aTea  out  their  true 
inip::aa:;:a  >.-  ^aa:  a  ;a-  a::::  ::  ra^  ::ra:^ra::a  :f  the  blood.  It 
was  a::  =;:  a\a:a  v;:-,;:  Ha:"T7  .rarziaa  irar;;  ra';a:;aas.  SmS  the  sttnir 
ulu=  ::  aii  ::ara:aaia  :aa:  aa:"'ea  ::  £a;a  ^'aa:  :;i::£:ajice  to  him, 
for  va  ;;-  ;^-  -  --  r.  ra^  ia^TaaiT  :a::r:  a:^  ar"  :i  the  purpose  of 
the  v&a.--=  0:  :::r  vria^  "■>-.  -zr:::^:;  a-.::rra::, 

In  1602.  Harvey  vras  gTaauatra  ^I.  D,  frz-ra  Paaaa.  His  diploma 
eonfarari  upon  him  the  degree  ::  L'lctar  of  Paa.a^a:.  "^iah  leave  to 
prac:::e  aad  teach  arts  and  mediciae  :a.  evei7"  land  ana  sea:  of  learn- 
ing^, I:  farther  stated,  that  "he  had  conducted  hianseh"  5:  ~;nder- 
fuii7  ~ef.  in  the  examination  anf  haf  shovm  saifa  .naaii,  anean:-7  and 

his  enaanners  had  formed  ai  him,  Ihe7  aenfrf.  "rneaeiire.  tnat  ne 
was  sahnah  e:>apert  and  most  eiai nenta.'  aaaanea  a:th  m  aaa=  and 
mecicme,  ana  to  this  the7  tut  tnen'  nan  is  anaa:n:nasl"a  ~a_nn.r.7 
and  ~i:h  complete  agreement  ana  aahrs:tatnm!7  "  The  Vnn.'-sasit:' 
of  Cambridge  conieiTed  the  degree  m  id,  I',  "an  naa:  :he  saane  7e'ta 

HarTe7  manied  in  1604.  the  daaghter  of  L'r,  Bai^mr.  ""n;  -.-aas 
ph7si:aan  to  Queen  Elizabeth  and  to  James  I. 

Haa-ve7.    as    ^e    shall    see,    excelled    as    lecttirer.      His    le:tur-s 

more  than  sixty  kinds  of  animals,  as  veil  as  a  thaatagh  ann"asfre  :f 
human  anatomy,  vrhich  must  ha'm  taaru  years  ■::  stafy  t:  anann. 
He  ~as  elected  feUow  of  the  Cillege  :f  Physicians  aa  '-"H'  _  --'--  --'-'-- 
portant  position  which  Hanrey  hela  '""as  Physnlaa  t:  St.  naatail;- 
mew's  Hospital  in  1609,  "The  charr^  :f  thr  rav-lnan  m  St  rar- 
tholome~'s  Hospital"  remlaea  tae  nmamhrnt  t:  aevme  at  leas;  n:e 
day  a  "nek  throughn^t  tne  :naa  t:  nmaaty.  Ha  ":"n.s  zamher  en;:lnen 
'"no":  t'.':  ic:'\'y..  l;nn.  n  azaa,  t:  aaanat  :a  "^"aate  i::ii~r!JJig  lor  tne 
poor  hat  sans  atml  aaf  '"hn-szme  talngs  as  he  snah  think  with  his 
oest  r       nr       1      a    a^     :n  am  tl ma:  aay  anection  or  respect  to 

be  naa  ta  tne  apothecary.  Ana  hr  ntall  tahe  no  gift  or  reward  of 
any  of  the  poor  of  this  house  for  his  counsel."  This  ''"'charge""  Har- 
vey is  said  to  have  faithfully  obsera-ed, 

Anatomical  Teaching  Previous  to  1743 — Duidng  Haimey's  day 
and  until  1745,  the  teaching  of  Anatomy  in  England  was  vested  in  a 
few  corporate  bodies.    Private  teaching  was  discouraged  by  fine  and 


WILLIAM  HARVEY  5 

imprisonment.  The  College  of  Physicians  and  Barber  Surgeons  had  a 
monopoly  in  London.  The  value  of  Anatomy  as  a  foundation  to  medi- 
cine was  fully  recognized  at  the  time  .  The  subjects  for  dissection 
were  the  bodies  of  executed  criminals.  Those  were  the  times  of 
public  executions,  witnessed  by  immense  crowds  whose  opposition 
and  sympathy  for  the  felon  and  his  friends  often  interfered  with  the 
procuring  of  the  body  for  dissection. 

The  method  of  anatomical  instruction  is  of  interest.  The  sub- 
ject was  taught  practically  by  a  series  of  demonstrations  on  the  body. 
The  absence  of  means  of  preservation  of  cadavers  precluded  instruc- 
tion in  detail.  A  single  body  was  dissected  to  show  the  muscles ;  an- 
other to  demonstrate  the  bones,  and  a  third  to  exhibit  the  viscera. 
Attendance  on  anatomical  lectures  and  demonstrations  was  com- 
pulsory; violation  meant  the  forfeiture  of  a  fine.  Some  were  ex- 
empted from  the  penalty,  as  one  entry  shows  that  a  Robert  Mudsley 
"has  licence  to  be  absent  from  all  lectures  without  payment  of  any 
fine,  because  he  has  given  over  the  art  of  surgery,  and  doth  occupy 
only  a  silk  shop  and  shave." 

The  anatomical  demonstrations  were  open  to  the  public.  The 
following  note  appears  in  Pepy's  Diary:*    "Up  and  to  my  office. 

.  .  .  Commissioner  Pett  and  I  walked  to  Chyrurgeon's  Hall  (we 
being  all  invited  thither,  and  promised  to  dine  there) ,  where  we  were 
led  into  the  Theater;  and  by  and  by  comes  the  reader.  Dr.  Teame^ 
with  the  master  and  company  in  a  very  handsome  manner;  and  all 
being  settled,  he  began  his  lecture,  this  being  the  second  upon  the 
ureters  and  kidneys,  which  was  very  fine;  and  his  discourse  being 
ended,  we  walked  into  the  hall,  and  there  being  a  great  store  of  com- 
pany, we  had  a  fine  dinner  and  good  learned  company,  many  Doctors 
of  Physique,  and  we  used  with  extraordinary  gi'eat  respect  .  .  .  After 
dinner  Dr.  Scarborough  took  some  of  his  friends,  and  I  went  along 
with  them  to  see  the  body  alone,  which  we  did,  which  was  a  lusty 
fellow,  a  seaman  that  was  hanged  for  a  robbery.  I  did  touch  the 
dead  body  with  my  bare  hand ;  it  felt  cold,  but  methought  it  was  a 
very  unpleasant  sight.  .  .  .  Thence  we  went  into  a  private 
room  where  I  perceive  they  prepare  the  bodies,  and  there  were  the 
kidneys  and  ureters,  etc.,  upon  which  he  read  today,  and  the  doctor, 
upon  my  desire  and  the  company's,  did  show  very  clearly  the  man- 
ner of  the  disease  of  the  stone  and  the  cutting  and  all  other  questions 
that  I  could  think  of."  Pepy's  interest  in  the  operation  of  cutting 
for  stone  is  said  to  be  due  to  the  fact  that  he  had  undergone  the 
ordeal  himself.  The  Dr.  Scarborough  mentioned  in  Pepy's  note  was 
a  friend  and  pupil  of  Harvey. 

Personal  Characteristics — Harvey  is  described  as  a  man  of  the 
"lowest  stature,  round  faced,  ^^ith  a  complexion  like  the  wainscot; 
his  eyes  small,  round,  very  black  and  full  of  spirit,  his  hair  black  as 
a  raven  and  curling ;  rapid  in  his  utterance,  chivalric  even  to  gesture, 
and  used  when  in  discourse  with  anyone  to  play  unconsciously  with 


♦Samuel  Pepys  (1632-1703),  was  a  famous  diarist.  His  Diary,  which 
extends  from  1660  to  1669,  was  written  in  shorthand,  and  was  deciphered 
by  Lord  Braybrooke  in  18  25.  This  delightful  book  of  gossip  is  one  of  the 
most  interesting  memorials  of  the  domestic  life  of  the  time. 


6  PATHFINDERS  OF  PHYSIOLOGY 

the  small  dagger  he  wore  by  his  side."  His  individuality  was  marked, 
as  was  evidenced  by  the  strong  impression  he  made  upon  those  with 
whom  he  came  in  contact.  His  intellectual  power  and  independence 
of  character  were  unusual.  His  interests  were  wider  than  his  scien- 
tific studies.  According  to  an  anonymous  biographer*  of  the  eight- 
eenth century,  "He  was  well  read  in  ancient  and  modem  history; 
and  when  he  was  wearied  with  too  close  attention  to  the  study  of 
nature,  he  would  relax  his  mind  by  discoursing  to  his  friends  on 
political  subjects  and  the  state  of  public  affairs.  He  took  great 
pleasure  in  reading  from  the  ancient  poets,  and  especially  Virgil, 
with  whose  work  he  was  exceedingly  delighted.  He  was  laboriously 
studious,  regular  and  virtuous  in  his  life  and  had  a  strong  sense  of 
religion.  In  his  familiar  conversation  there  was  a  mixture  of  gravity 
and  cheerfulness;  he  expressed  himself  with  great  perspicuity,  and 
with  much  grace  and  dignity;  and  was  eminent  for  his  great  candor 
and  moderation.  He  never  endeavored  to  detract  from  the  merit  of 
other  men;  but  appeared  always  to  think  that  the  virtues  of  others 
were  to  be  imitated  and  not  envied." 

In  spite  of  his  choleric  and  hasty  disposition  he  had  the  faculty 
of  making  close  friendships.  His  replies  to  his  critics  showed  great 
moderation.  Harvey's  true  character  is  probably  best  seen  in  that 
period  of  his  life  which  was  beset  with  opposition  and  reproach,  im- 
mediately following  the  publication  of  his  great  work  on  the  circu- 
lation. To  his  traducers  his  attitude  resembled  that  of  the  divine 
Master,  "To  return  evil  speaking  with  evil  speaking  I  hold  to  be  un- 
worthy of  a  philosopher  and  searcher  after  truth.  I  believe  I  shall 
do  better  and  more  advisedly  if  I  meet  so  many  indications  of  ill- 
breeding  with  the  light  of  faithful  and  conclusive  observation."  His 
attitude  also  resembles  that  of  Darwin  who,  on  the  publication  of  his 
Origin  of  the  Species,  was  met  with  a  storm  of  abuse  from  clerical 
ignorance.  It  is  said  that  the  great  evolutionist  not  only  observed  a 
tranquility  impassionate  and  unique  but  even  condescended  to  reply  at 
length  with  courtesy  to  the  rantings  of  those  who  vilified  without 
even  reading  his  work  or  comprehending  the  object  of  their  denunci- 
ations. 

Harvey  was  not  a  religious  man  in  the  narrow  sense  of  the  term 
despite  the  fact  that  he  lived  in  an  age  of  warring  creeds.  His  views 
were  broad  as  befitted  a  student  of  the  design  and  workmanship  of 
the  Great  Architect  of  the  universe.  According  to  Sir  Russell  Rey- 
nolds, "a  devout  and  reverential  recognition  of  God"  permeated  his 
work,  "not  only  as  the  great  primal  ever-acting  force,  defined  outside 
and  before  all  the  works  of  nature;  but  as  the  Being,  'the  Almighty 
and  Eternal  God'  to  whom  he  says  in  his  last  will  and  testament,  *I 
do  most  humbly  render  my  soul  to  Him  who  gave  it;  and  to  my 
blessed  Lord  and  Saviour  Jesus  Christ.'  " 

Harvey's  knowledge  of  Latin  was  so  thorough  that  he  could  con- 
verse with  facility  equal  to  his  native  tongue.  He  was  accustomed 
to  employ  both  English  and  Latin  even  in  the  same  sentence,  for  ex- 
ample, speaking  of  the  eyes  and  their  function:  "Oculi  eodem  loco, 
viz,  nobilissimi  supra  et  ante  ad  processus  eminentes  instar  capitis 
in  a  lobster  snayles  cornubus  tactu  pro  visu  utuntur  unde  oculi  as  a 
centinell  to  the  army  locis  editis  anterioribus." 

♦British   Biographies,   Vol.    IV.,   London,    1768. 


WILLIAM  HARVEY  7 

Harvey  as  Lecturer — Harvey's  lectures  were  partly  read  and 
partly  oral.  The  cadaver  lay  on  the  table  with  the  dissecting  instru- 
ments close  to  it.  An  assistant  dissected  or  demonstrated  while  the 
lecturer  read  his  remarks.  The  anatomical  lecturer  of  the  sixteenth 
century  was  a  personage  of  importance.  The  greatest  consideration 
was  exercised  for  his  personal  comfort.  The  stewards  were  instruct- 
ed, "to  see  and  to  provide  that  there  be  a  mat  about  the  hearth  in  the 
hall  that  the  Doctor  be  made  not  to  take  cold  upon  his  feet.  *  *  * 
And  further,  that  there  be  two  fine  white  rods  appointed  for  the 
Doctor  to  touch  the  body  where  it  shall  please  him ;  and  a  wax  candle 
to  look  into  the  body,  and  that  there  be  always  for  the  Doctor  two 
aprons  to  be  from  the  shoulder  dov^Tiward  and  two  pair  of  sleeves 
for  his  whole  arm.  .  .  .  and  not  to  occupy  one  apron  and  one 
pair  of  sleeves  every  day,  which  is  unseemly."  Harvey  laid  down  the 
following  precepts  for  his  own  guidance  as  lecture  precepts  which 
the  modem  anatomical  lecturer  might  observ^e  with  propriety: 

(1)  To  show  as  much  as  may  be  at  a  glance,  the  whole  belly 
for  instance,  and  afterwards  to  subdivide  the  parts  according  to  their 
position  and  relations. 

(2)  To  point  out  what  is  peculiar  to  the  actual  body  being  dis- 
sected. 

(3)  To  supply  only  by  speech  what  cannot  be  shown  on  your 
own  credit  and  authority. 

(4)  To  cut  up  as  much  as  may  be  in  the  sight  of  the  audience. 

(5)  To  enforce  the  right  opinion  by  remarks  down  from  far 
and  near  and  to  illustrate  more  by  the  structure  of  animals  accord- 
ing to  the  Socratic  rule. 

(6)  Not  to  praise  or  dispraise  other  anatomists,  for  all  did  well 
and  there  was  some  excuse  even  for  those  who  are  in  error. 

(7)  Not  to  dispute  with  others. 

(8)  To  state  things  briefly  and  plainly, 

(9)  Not  to  speak  of  anything  which  can  be  explained  without 
the  body  or  can  be  read  at  home. 

Here  we  have  a  combination  of  orthodox  medical  ethics  and 
sound  pedagogy.  Harvey's  particular  role  as  *Lumlian  lecturer  in- 
cluded the  position  of  lecturer  upon  the  viscera.  Discussing  the  tho- 
racic viscera  he  ennunciated  the  remarkable  discovery  with  which 
his  name  is  inseparably  associated,  initialing  the  notes  to  indicate 
that  the  ideas  were  peculiarly  his  own. 

constat  per  fabricam  cordis  sanguinem. 

per  pulmones  in  Aortam  perpetuo. 

Transf  erri,  as  by  two  clacks  of  a 

water  bellows  to  rayse  water. 

constat  per  ligaturam  transitum  saguinis 

ab  arteriis  ad  venas 

unde  perpetuum  sanguinis  motum 

in  circulo  fieri  pulsu  cordis. 

W.  H. 

*Tlie  Lumlian  lecture  was  a  surgical  lecture  established  at  a  cost  of 
£40  a  year,  which  sum  accrued  from  the  rental  of  lands  of  Lord  Lumlej,  of 
Essex,  England. 


8  PATHFINDERS  OF  PHYSIOLOGY 

"It  is  plain  from  the  structure  of  the  heart  that  the  blood  is 
passed  continuously  through  the  lungs  to  the  aorta  as  by  the  two 
clacks  of  a  water  bellows  to  raise  water. 

"It  is  shown  by  the  application  of  a  ligature  that  the  passage  of 
the  blood  is  from  the  arteries  into  the  veins. 

"Whence  it  follows  that  the  movement  of  the  blood  is  constantly 
in  a  circle  and  is  brought  about  by  the  beat  of  the  heart."  It  was 
not  until  twelve  years  after  this  important  announcement  that  he 
proclaimed  it  to  a  wider  audience. 

Harvey's  literary  style  was  somewhat  figurative.  He  loved  to 
indulge  in  metaphors — witness: 

An  cerebrum  rex,  whether  the  brain  is  king. 

Nervi  majistratus,  the  nerves  his  ministers. 

Musculi  cives  populus,  the  muscles  the,  citizens  or  the  people. 

He  also  draws  a  similtude  liking  the  brain  to  a  military  com- 
mander, the  leader  of  an  orchestra,  an  architect,  and  he  speaks  of  the 
muscles  and  nerves  as  subordinate  officers. 

Year  by  year  Harvey  delivered  the  Lumlian  lectures  to  the  Col- 
lege of  Physicians.  His  private  practice  grew  so  as  to  be  fairly  lucra- 
tive. 

Harvey  and  Bacon — In  1618  he  was  appointed  physician  to 
James  I.  In  1631  he  was  appointed  physician  in  ordinary  to  King 
James'  son,  Charles  I.  Not  only  gained  he  an  entrance  to  the  house- 
hold of  the  king  but  he  was  employed  in  the  homes  of  the  most  dis- 
tinguished nobles.  Among  others  he  attended  Sir  Francis  Bacon, 
who  was  always  a  weak  and  ailing  man  with  a  disposition  to  be  hypo- 
chondriac. "In  William  Harvey  and  Francis  Bacon,"  says  Gorton, 
may  be  observed  two  men  Hke  planets  in  conjunction;  bom  in  the 
same  generation,  each  illustrious  in  the  annals  of  history,  the  one  in 
philosophy,  the  other  in  science  but  in  striking  contrast  to  each 
other.  The  one  was  a  thinker,  the  other  was  an  actor;  one  con- 
ceived methods,  the  other  put  methods  into  operation;  one  was  an 
academic  philosopher,  the  other  a  man  of  science  and  discovery;  one 
immortalized  himself  by  his  profundity  of  thought,  the  other  by  his 
contribution  to  science.  Both  were  stars  in  the  firmament  of  great 
men,  but  long  after  one  has  become  dim  or  gone  out,  the  other  will 
continue  to  shine  with  splendor." 

Though  honored  by  England's  Lord  Chancellor  as  the  custodian 
of  his  health,  Harvey  evidently  failed  to  be  impressed  with  Bacon's 
greatness  even  as  philosopher,  for  speaking  of  him,  Harvey  refers  to 
him  as  "writing  philosophy  like  a  Lord  Chancellor." 

Publication  of  His  Work  on  the  Circulation — In  1628,  the  crown- 
ing event  of  his  life  took  place  when  he  published  his  well  considered 
and  matured  account  of  the  circulation  of  the  blood.  He  had  demon- 
strated his  ideas  of  the  circulation  for  twelve  years  before  publishing 
them,  which  event  occurred  in  the  fiftieth  year  of  his  life.  This 
monumental  work  of  the  great  physiologist  was  accomplished  while  yet 
in  his  thirties.  Why  Harvey  should  allow  so  much  time  to  elapse  be- 
tween the  event  of  his  epochal  discovery  and  its  publication  is  not 
clear.  Evidently  the  passion  to  rush  into  print  was  not  so  great  as 
it  is  with  the  investigator  of  to  day.    It  is  interesting  to  note,  how- 


WILLIAM  HARVEY  9 

ever,  that  among  the  greatest  thinkers  and  investigators  Harvey  is 
not  unique  in  this  respect.  Copernicus  is  said  to  have  detained  his 
"Treatise  of  Revolutions"  thirty  years  before  permitting  its  publi- 
cation; Bacon  kept  his  Novum  Organum  by  him  for  twelve  years; 
Isaac  Newton  ''brooded  in  silence  over  the  motion  of  the  spheres" 
for  more  than  twenty  years  before  pubhshing  his  Principia ;  between 
the  first  draft  and  the  publication  of  the  Origin  of  the  Species  seven- 
teen years  were  permitted  to  intervene.  Perhaps  it  was  Harvey's 
reluctance  toward  "quitting  the  peaceful  haven,"  that  constrained 
him  for  so  long  a  time,  for  elsewhere  he  tells  us  that  his  practice 
fell  off  or,  to  use  his  own  words,  he  "fell  mighty  in  practice."  Re- 
garding him  a  contemporary  wrote,  "though  all  of  his  profession 
would  allow  him  to  be  an  excellent  anatomist,  I  never  heard  of  any 
who  admired  his  therapeutic  way.  I  knew  several  practitioners  in 
this  town  that  would  not  have  given  three  pence  for  his  bills  (pre- 
scriptions) as  a  man  can  hardly  tell  by  his  bills  what  he  did  aim  at." 
Harvey  is  said  to  have  been  the  first  to  be  persecuted  by  the  medical 
profession  for  making  discoveries  at  variance  with  the  drift  of  public 
thought  and  opinion.  The  story  of  all  discoveries  of  the  first  rank 
has  borne  out  Locke's  aphorism  that  "Truth  scarce  ever  yet  carried 
by  vote  at  its  first  appearance."  The  greatest  obstacle  to  the  accept- 
ance of  truth  seems  to  be  our  present  knowledge.  Men  are  by  nature 
conservative;  they  resent  innovations.  Bagehot  tells  us  that  the 
"pain  of  a  new  idea  is  one  of  the  greatest  pains  to  human  nature." 
Socrates  somewhere  likens  himself  to  a  midwife  but  his  peculiar 
function  in  life  was  to  assist  in  that  mental  labor  which  gave  birth 
to  ideas,  a  similitude  which  is  suggestive  of  pain.  The  man  who  ex- 
presses a  new  idea  is  apt  to  be  abused,  perhaps  stoned.  ¥/hatever 
may  be  said  of  the  twentieth  century  the  scientific  world  can  be  ac- 
cused no  longer  of  tardiness  in  the  acceptance  of  new  truth,  but  it 
reserves  the  right  to  "prove  all  things  and  to  hold  fast  to  that  which 
is  good."  While  Harvey's  practice  may  have  fallen  off,  his  discovery 
did  not  by  any  means  consign  him  to  obscurity.  He  still  found  favor 
with  King  Charles  I,  whose  personal  physician  he  was.  His  constant 
attendance  at  court  greatly  interfered  with  his  duties  at  St  .Bar- 
tholomew's Hospital  and  resulted  in  the  appointment  of  an  assistant, 
but  with  no  diminution  in  Harvey's  stipend.  A  contemporary  of  Har- 
vey states  as  follows:  "I  have  heard  him  say  that  after  his  Booke 
of  Circulation  of  the  Blood  came  out  he  fell  mightily  in  practice,  and 
'twas  believed  by  the  vulgar  that  he  was  crack-brained,  and  all  the 
physicians  were  against  him,  with  much  adoe  at  last  in  about  twenty 
or  thirty  years'  time  it  was  received  in  all  the  universities  of  the 
world,  and  as  Dr.  Hobbs  says  in  his  book  'De  Corpore,'  he  is  the  only 
man  perhaps  that  ever  lived  to  see  his  own  doctrine  established  in  his 
lifetime;"    v^eritas  est  magna  et  prevalebit! 

And  yet,  after  the  discovery  has  been  recognized  as  one  of  mo- 
mentous import,  the  scientist  has  his  detractors.  Harvey  was  no  ex- 
ception. There  were  those  who  sought  to  disprove  the  originality  of 
his  work.  Some  attributed  the  merit  of  discovering  the  circula- 
tion to  Servetus,  some  to  Realdus  Columbus,  others  to  Caesalpinus. 
True,  Servetus,  a  Spaniard,  bom  in  1511  and  burned  at  the  stake  in 
Geneva,  1533,  at  the  bidding  of  Calvin,  in  a  copy  of  his  Restitutio, 
which  was  saved  when  an  edition  of  1,000  copies  met  the  fate  of  the 
author,  rejected  the  contention  that  the  blood  passed  through  the 


10  PATHFINDERS  OF  PHYSIOLOGY 

cardiac  septum.  He  had  grasped  the  true  features  of  the  pulmonary 
circulation — the  passage  of  the  blood  from  the  right  side  to  the 
lungs,  thence  to  the  left  side  or  ventricle.  Realdus  Columbus,  born 
at  Cremona,  1516,  a  presumptuous  personage,  speaks  of  the  blood 
carried,  "by  the  artery-like  vein  to  the  lung  and  being  there  made 
thin  is  brought  back  thence  together  with  air  by  the  vein-like  artery 
to  the  left  ventricle  of  the  heart."  Then  he  goes  on  to  press  his  claim 
by  declaring  that,  hitherto,  no  one  had  made  this  observation  or  re- 
corded it  in  writing.  Andreas  Caesalpinus  was  bom  at  Arezzo  in 
1519.  He  held  for  many  years  the  professorship  of  medicine  at  Pisa. 
Learned  in  all  the  lore  of  the  ancients,  he  was  noted  among  other 
things  for  his  determined  opposition  to  Galen;  Caesalpinus  appears 
to  have  grasped  one  important  truth,  namely,  that  the  heart  at  systole 
discharges  its  contents  into  the  aorta  and  pulmonary  artery,  and  at 
its  diastole  receives  blood  from  the  vena  cava  and  pulmonary  vein. 
Let  all  this  be  granted,  yet  the  great  work  of  Harvey  is  not  a 
whit  less  meritorious.  The  steam  engine  was  in  existence  before  the 
day  of  James  Watt,  yet  his  name  is  inseparably  associated  with  the 
invention  which  transformed  a  mere  toy  into  a  gigantic  factor  which 
has  revolutionized  human  industry.  No  person,  not  even  the  genius  is 
independent  of  his  time ;  he  is  the  heir  of  all  the  ages,  and  his  great- 
ness does  not  depend  so  much  in  presenting  something  unprecedented 
as  it  does  in  seeing  something  clearly  and  telling  in  a  simple  way 
what  he  has  seen. 

Treatise  on  the  Circulation. — Harvey's  greatest  work  was  un- 
doubtedly his  Exercitatio  Anatomica  de  Motu  Cordis  et  Sanguinis  in 
Animalibus,  an  anatomical  treatise  on  the  movement  of  the  heart 
and  blood  in  animals,  published  in  Frankfort,  Germany,  in  1628.  The 
book  was  a  small  quarto  volume  of  72  pages.  It  opens  with  a  dedi- 
cation to  "The  Most  Illustrious  and  Indomitable  Prince,  Charles,  King 
of  Great  Britain,  France,  and  Ireland,  Defender  of  the  Faith,"  etc. 
The  dedication  proceeds :  "The  heart  of  animals  is'  the  foundation  of 
their  life,  the  sovereign  of  everything  within  them,  the  sun  of  their 
microcosm,  that  upon  which  all  growth  depends,  from  which  all 
power  proceeds.  The  king  in  like  manner,  is  the  foundation  of  his 
kingdom,  the  sun  of  the  world  around  him,  the  heart  of  the  republic, 
the  fountain  whence  all  power,  all  grace  doth  flow."  Whatever  may  be 
said  regarding  Charles  I,  who  was  the  victim  of  public  execution,  he 
certainly  befriended  Harvey.  Then  to  the  president  of  the  Royal  Col- 
lege of  Physicians  and  to  other  learned  physicians  the  author  ad- 
dresses himself  in  a  dedication  which  he  concludes :  *  *  *  "I  profess 
both  to  learn  and  to  teach  anatomy  not  from  books  but  from  dissec- 
tions; not  from  the  positions  of  philosphers  but  from  the  fabric  of 
nature.  *  *  *  i  avow  myself  the  partisan  of  truth  alone;  and  I  can 
indeed  say  that  I  have  used  all  my  endeavors,  bestowed  all  my  pains 
on  an  attempt  to  produce  something  that  should  be  agreeable  to  the 
good,  profitable  to  the  learned,  and  useful  to  letters."  Harvey's 
method  here  ennunciated  is  the  method  of  every  scientist  since  his 
day,  whose  contribution  has  possessed  real  merit — that  is,  reasoning 
based  upon  experiment  and  observation. 

The  work  on  the  circulation  comprises  seventeen  short  chapters. 
It  is  an  interesting  account,  lucid  and  connected,  of  the  heart's  action 
and  the  circulation  of  the  blood.    Harvey  had  no  means  of  knowing 


WILLIAM  HARVEY  11 

the  connection  between  the  smallest  arteries  and  the  smallest  veins, 
for  the  microscope  was  not  in  such  a  stage  of  perfection  as  to 
permit  of  much  fine  work  in  minute  anatomy.  It  was  not  until  the 
invention  of  the  compound  microscope  in  1675  that  Leeuwenhoek 
described  blood  corpuscles  and  the  capillary  circulation.  In  the  first 
chapter  the  author  reviews  some  of  the  fantastic  theories  regarding 
the  functioning  of  heart  and  lungs.  The  heart  was  held  to  be  the 
great  heat  center  of  the  body.  The  blood  was  sucked  into  it  during 
diastole  and  expelled  from  it  during  systole.  The  arteries  cooled  the 
blood;  the  lungs  fanned  and  cooled  the  heart.  The  term  "spirits" 
meant  a  great  deal  to  Harvey's  predecessors,  but  not  to  him.  "The 
word  blood  has  nothing  of  grandiloquence  about  it,  for  it  signifies  a 
substance  which  we  have  before  our  eyes  and  can  touch;  but  before 
such  titles  as  spirit  and  calidum  innatum  (inherent  heat)  we  stand 
agape." 

Chapter  I,  he  continues: 

"When  I  first  gave  my  mind  to  vivisections,  as  a  means  of  discovering  tlie 
motions  and  uses  of  ttie  heart,  and  sought  to  discover  these  from  actual  inspec- 
tion, and  not  from  the  writings  of  others,  I  found  the  task  so  truly  arduous,  so 
full  of  difficulties,  that  I  was  almost  tempted  to  think,  with  Fracastorius,  that 
the  motion  of  the  heart  v/as  only  to  be  comprehended  by  God.  For  I  could 
neither  rightly  perceive  at  first  the  systole  and  when  the  diastole  took  place,  nor 
when  and  where  dilatation  and  contraction  occurred,  by  reason  of  the  rapidity 
of  the  motion,  which  in  many  animals  is  accomplished  in  the  twinkling  of  an  eye, 
coming  and  going  like  a  fiash  of  lightning;  so  that  the  systole  presented  itself  to 
me  now  from  this  point,  now  from  that;  the  diastole  the  same;  and  then  every- 
thing was  reversed,  the  motions  occurring,  as  it  seemed,  variously  and  confusedly 
together.  *  *  * 

"At  length,  and  by  using  greater  and  daily  diligence,  having  frequent  re- 
course to  vivisections,  employing  a  variety  of  animals  for  the  purpose,  and  collat- 
ing numerous  observations,  I  thought  that  I  had  attained  to  the  truth,  that  I 
should  extricate  myself  and  escape  from  this  labyrinth,  and  that  I  had  discovered 
what  I  so  much  desired,  both  the  motion  and  the  use  of  the  heart  and  arteries'; 
since  which  time  I  have  not  hesitated  to  expose  my  views  upon  these  subjects, 
not  only  in  private  to  my  friends  but  also  in  public,  in  my  anatomical  lectures 
after  the  manner  of  the  academy  of  old." 

He  goes  on  to  tell  how  his  views  pleased  some,  displeased  others. 

He  finds  it  advantageous  to  study  the  movement  of  the  heart  in 
the  cold-blooded  animals — frogs,  snakes  and  fishes.  He  ascertained 
that  the  heart  was  a  muscular  organ,  that  its  systole  was  the  result 
of  muscular  contraction.  The  contraction  of  the  heart  was  more  im- 
portant than  its  dilitation.  "During  its  contraction  the  heart  becomes 
erect,  hard  and  diminished  in  size,  so  that  the  ventricles  become 
smaller  and  are  so  made  more  apt  to  expel  their  charge  of  blood.  In- 
deed, if  the  ventricle  be  pierced  the  blood  will  be  projected  forcibly 
outward  at  each  pulsation  when  the  heart  is  tense."  Harvey  showed 
that  the  pulsation  of  the  arteries  depended  upon  the  contraction  of 
the  left  ventricle.    The  contraction  of  the  right  ventricle  propelled 

♦The  extracts  which  follow  illustrate  Harvey's  style.  The  Motion  of  the 
Heart  and  Blood,  by  William  Harvey,  can  be  procured  in  convenient  form 
In  the  Everyman's  Library  Series  (E.  P.  Dutton  &  Co.,  New  York).  This  is 
a    reprint   from   the   Sydenham   Society's  edition  of   1847. 


12  PATHFINDERS  OP  PHYSIOLOGY 

the  blood  into  the  pulmonary  arteries,  the  pulsations  of  which  were 
simultaneous  with  the  other  arteries  of  the  body.  He  demonstrated 
that  the  two  ventricles  contracted  simultaneously  and  that  the  two 
auricles  contracted  at  the  same  time. 

Motion,  Action  and  Office  of  the  Heart. — In  the  fifth  chapter 
Harvey  deals  with  the  motion  and  function  of  the  heart.  It  readi 
somewhat  like  a  modem  work  in  physiology. 

"First  of  all,  the  auricle  contracts,  and  in  the  course  of  its  contraction 
throws  the  blood  (which  it  contains  in  ample  quantity  as  the  head  of  the  veins, 
the  storehouse,  and  cistern  of  the  blood),  into  the  ventricle,  which,  being  filled, 
the  heart  raises  itself  straightway,  makes  all  its  fibres  tense,  contracts  the  ven- 
tricles, and  performs  a  beat,  by  which  beat  it  immediately  sends  the  blood  sup- 
plied to  it  by  the  auricle  into  the  arteries;  the  right  ventricle  sending  its  charge 
into  the  lungs  by  the  vessel  which  is  called  vena-arteriosa,  but  which,  in  structure 
and  function,  and  all  things  else,  is  an  artery;  the  left  ventricle  sending  its 
charge  into  the  aorta,  and  through  this  by  the  arteries  to  the  body  at  large. 
These  two  motions,  one  of  the  ventricles,  another  of  the  auricles,  take  place  con- 
secutively, but  in  such  a  manner  that  there  is  a  kiud  of  harmony  or  rhythm  pre- 
served between  them,  the  two  concurring  in  such  wise  that  but  one  motion  is 
apparent,  especially  in  the  warmer  blooded  animals,  in  which  the  movements  in 
question  are  rapid." 

So  far  as  Harvey's  reasoning  is  based  upon  his  observations  his 
conclusions  are  in  the  main  correct,  as  proved  by  more  recent  re- 
search ;  where  he  indulges  in  speculation  we  get  the  following : 

"In  the  larger  and  more  perfect  animals  of  mature  age  Nature 
has  rather  chosen  to  make  the  blood  percolate  the  parenchyma  of  the 
lungs.  *  *  *  It  must  be  because  the  larger  and  more  perfect  animals 
are  warmer,  and  when  adult  their  heat  greater,  ignited  I  may  say 
and  requiring  to  be  damped  or  mitigated,  that  the  blood  is  sent 
through  the  lungs,  in  order  that  it  may  be  tempered  by  the  air  that 
is  inspired  and  prevented  from  boiling  up  and  so  becoming  extin- 
guished or  something  else  of  the  sort,"  or,  to  modernize  it,  the  lungs 
serve  as  radiator  and  the  heart  the  gasoline  or  internal  combustion 
engine. 

Capillary  Circulation. — Since  Harvey's  time  Malpighi,  in  1661, 
hinted  at  the  capillary  circulation,  which  was  still  further  investi- 
gated by  Leuwenhoek  in  1674,  who  studied  it  with  his  microscope  in 
the  web  of  a  frog's  foot  and  in  other  transparent  membranes.  In 
1676,  Blankaart,  and  in  1697  Cowper,  studied  the  arrangement  of  the 
capillaries  by  means  of  injected  specimens.  A  long  interval  elapsed 
between  the  histological  study  of  the  circulation  before  chemistry 
was  sufficiently  advanced  to  afford  definite  knowledge  in  regard  to 
oxidation  of  the  blood  and  the  explanation  of  the  true  function  of 
the  lungs.  The  work  of  Priestly  in  1775  was  a  notable  contribution 
to  the  physiology  of  respiration.  The  nineteenth  century,  through 
the  work  of  Ludwig  in  Germany,  Chauveau  in  France,  and  Foster  in 
England,  has  seen  the  physics  of  the  heart  and  circulation  reduced 
almost  to  an  exact  science. 

Any  account  of  the  works  of  Harvey  would  be  incomplete  were 
no  mention  made  of  his  work  in  embryology.  Harvey  discussed  the 
nature  of  development  and  exhibited  extraordinary  powers  as  re- 


WILLIAM  HARVEY  13 

gards  accuracy  of  reasoning-.  He  may  be  considered  as  having  made 
the  first  independent  advance  in  the  subject.  That  he  did  not  ac- 
compHsh  more  was  due  to  lack  of  instruments  of  precision,  and  to 
the  fact  that  he  had  to  build  on  the  general  level  of  the  science  of 
the  time.  His  work  on  embryology  was  published  in  1651.  It  was 
entitled  "Exercitationes  de  Generatione  Animalium."  In  it  is  an 
account  of  not  only  the  development  of  the  chick,  but  of  deer  and 
other  mammals  as  well. 

All  honor  to  him  who  blazes  the  trail.  The  refinements,  what- 
ever they  may  be,  can  never  merit  for  the  investigator  the  honor 
which  is  due  the  pioneer.  As  was  said  by  Haller,  one  of  the  best 
informed  minds  of  the  eighteenth  century,  "It  is  not  to  Caesalpinus, 
because  of  some  words  of  doubtful  meaning,  but  to  Harvey,  the  able 
writer,  the  laborious  contriver  of  so  many  experiments,  the  staid 
propounder  of  all  the  arguments  available  in  his  day,  that  the  im- 
mortal glory  of  having  discovered  the  circulation  of  the  blood  is  to 
be  assigned." 

One  of  his  last  acts  was  to  set  aside  a  certain  sum  derived  from 
his  estate  for  the  delivery  of  an  oration  in  commemoration  of  the 
benefactors  of  the  College  of  Physicians.  This  oration,  the  Har- 
veian  Oration,  is  still  delivered  each  year  by  some  distinguished  mem- 
ber of  the  medical  profession.  Even  in  his  declining  years  his 
thoughts  were  turned  to  the  future.  The  Harveian  Lecture  is  in- 
tended to  further  the  progress  of  science,  especially  a  knowledge  of 
the  body  in  health  and  disease.  "Much  of  the  nobility  of  the  profes- 
sion," says  Osier,  Harveian  lecturer,  1906,  "depends  upon  the  great 
cloud  of  witnesses'  who  pass  into  the  silent  land — pass  and  leave  no 
sign,  becoming  as  though  they  had  never  been  bom.  And  it  was  the 
pathos  of  this  fate  not  less  prophetic  because  common  to  all  but  the 
few,  that  wrung  from  the  poet  that  sadly  true  comparison  of  the  race 
of  man  to  the  race  of  the  leaves."  Harvey  was  one  of  the  "few"  to 
have  achieved  that  immortality  which  places  him  with  "The  divine 
men  of  old  time." 

He  died  June  3rd,  1657,  in  the  eightieth  year  of  his  age. 

AseUius  and  the  Lymphatic  Circulation. 

Corollary  to  the  circulation  of  the  blood  is  the  lymphatic  circula- 
tion. The  discovery  of  the  lymphatics  was  almost  synchronous  with 
that  with  which  Harvey  achieved  an  immortal  name.  While  the 
memory  of  Harvey  has  been  fittingly  honored  in  various  ways,  that 
of  Asellius  or  Aselli  has  not  been  sufficiently  recognized.  The  data 
referring  to  Aselli's  life  are  extremely  meagre.  He  was  bom  in  1581, 
at  Cremona,  Italy,  the  descendant  of  a  patrician  family.  He  studied 
at  the  University  of  Pavia,  where  he  became  laureate  in  medicine, 
surgery  and  philosophy,  after  which  he  located  in  Milan,  where  he 
taught  anatomy  privatoly  and  engaged  in  the  practice  of  surgery.  It 
was  while  in  Milan  that  he  made  his  discovery,  in  1622,  of  the  lym- 
phatic vessels  which  he  called  venae  lactae.  His  discovery  was  rec- 
ognized by  his  election,  two  years  later,  to  the  chair  of  anatomy  and 
surgery  in  his  alma  mater,  a  position  he  was  destined  not  long  to 
hold,  for  he  died  in  1626  at  the  age  of  forty-five.  His  book  De  Lac- 
tibus  was  published  a  year  after  his  death.     William  Harvey  was 


14  PATHFINDERS  OF  PHYSIOLOGY 

forty-four  years  old  at  the  time  of  Aselli's  discovery.     Aselli's  dis- 
covery of  the  lacteals  is  related  by  himself  as  follows : 

"On  the  23rd  of  July  in  thati  year  (1622)  I  had  taken  a  dog  in  good  condi- 
tion and  well  fed,  for  a  vivisection  at  tlie  request  of  some  friends,  wlio  very 
much  wished  to  see  the  recurrent  nerves.  When  I  had  finished  this  demonstra- 
tion of  the  nerves,  it  seemed  good  to  watch  the  movements  of  the  diaphragm  in 
the  same  dog,  at  the  same  operation.  While  I  was  attempting  this,  and  for  that 
purpose  had  opened  the  abdomen  and  was  pulling  down  with  my  hand  the  intes- 
tines and  stomach  gathered  together  into  a  mass,  I  suddenly  beheld  a  great 
number  of  cords,  as  it  were,  exceedingly  thin  and  beautifully  white,  scattered 
all  over  the  whole  of  the  mesentery  and  the  intestine,  and  starting  from  almost 
innumerable  beginnings.  At  first  I  did  not  delay,  thinking  them  to  be  nerves. 
But  presently  I  saw  I  was  mistaken  in  this,  since  I  noticed  the  nerves  belonging 
to  the  intestine  were  distinct  from  these  cords  and  wholly  unlike  them,  and, 
besides,  were  distributed  quite  separately  from  them.  Wherefore  struck  by  the 
novelty  of  the  thing,  I  stood  for  some  time  silent  while  there  came  to  my  mind 
the  various  disputes,  rich  in  personal  quarrels  no  less  than  in  words,  taking  place 
among  anatomists  concerning  the  mesariac  veins  and  their  function.  And  by 
chance  it  happened  that  a  few  days  before  I  had  looked  into  a  little  book  by 
Johannes  Costaeus  written  about  this  very  matter.  When  I  gathered  my  wits 
together  for  the  sake  of  the  experiment,  having  laid  hold  of  a  very  sharp  scalpel, 
I  pricked  one  of  those  cords,  and  indeed  one  of  the  largest  of  them.  I  had 
scarcely  touched  it,  when  I  saw  a  white  liquid  like  milk  or  cream  forthwish  gush 
out.  Seeing  this,  I  could  hardly  restrain  niy  delight,  and  turning  to  those  who 
were  standing  by,  to  Alexander  Tadinus,  and  more  particularly  to  Senator  Sep- 
talius,  who  was  both  a  member  of  the  great  college  of  the  Order  of  Physicians 
and,  while  I  am  writing  this,  the  medical  officer  of  health,  'Eureka,'  I  exclaimed 
with  Archimedes,  and  at  tb.e  same  time  invited  them  to  the  interesting  spectacle 
of  such  an  unusual  phenomenon.  And  they  indeed  were  much  struck  with  the 
novelty  of  the  thing." 

Aselli  noted  the  presence  of  valves  in  the  lymphatic  vessels  and 
recognized  their  function,  namely,  to  prevent  the  backward  flow  of 
the  lymph.  He  recognized  also  that  the  lacteals  were  vessels  for  con- 
veying chyle  away  from  the  intestine.  He  went  wrong,  however,  in 
regard  to  the  ultimate  course  taken  by  the  newly-discovered  vessels, 
for  he  thought  he  could  trace  them  to  the  liver.  Aselli  was  heavily 
handicapped  by  his  previous  learning,  which  consisted  of  a  careful 
study  of  as  well  as  veneration  for  the  teachings  of  the  ancients. 
Galen  had,  in  fact,  taught  that  all  nutritive  material  from  digestive 
processes  passed  through  the  liver.  Aselli  speaks  in  his  book  of  a 
group  of  lymphatic  glands  lying  in  the  mesentery,  as  the  pancreas — 
hence  the  name  pancreas  Aselli.  The  force  which  caused  the  move- 
ment of  the  fluid  in  the  lacteal  vessels  was  believed  by  him  to  be  two- 
fold, a  vis  a  tergo  and  a  vis  a  f route;  the  latter  derived  from  supposed 
suction  of  the  liver  and  the  former  supplied  by  the  movements  of  the 
intestines. 

Aselli  Opposed  by  Harvey.  Aselli  in  his  modesty  endeavored  to 
prove  that  the  lacteals  were  known  to  the  ancients,  especially  to 
Herophilus  and  Erasistratus,  founders  of  the  Alexandrine  school  of 
medicine.  His  discovery  met  the  same  opposition  as  did  Harvey's,  and 
from  the  same  men,  among  them  Riolan  and  Primrose,  and  strange  to 
say  Harvey  himself  failed  to  recognize  the  importance  of  the  work 


ASELLIUS  15 

of  his  contemporary.     In  a  private  letter  written  in  April,  1652,  he 
writes : 

"With  regard  to  the  lacteal  veins  discovered  by  Aselli,  and  by  the  further 
diligence  ot!  Pecquet,  who  discovered  the  receptacle  or  reservoir  of  the  chyle, 
and  traced  the  canals  thence  to  the  sub-clavian  veins,  I  sTiall  tell  you  freely,  since 
you  ask  me,  what  I  think  of  them.  I  had  already,  in  the  course  of  my  dissections, 
I  venture  to  say  even  before  Aselli  had  published  hisi  book,  observed  these  white 
canals.  *  *  *  But,  for  various  reasons,  and  led  by  several  experiments,  I 
could  never  be  brought  to  believe  that  that  milky  fluid  was  chyle,  conducted  thither 
from  the  intestines,  and  distributed  to  all  parts  of  the  body  for  their  nourish- 
ment; but  that  it  was  rather  met  with  occasionally  and  by  accident,  and  proceeded 
from  too  ample  supply  of  nourishment  and  a  peculiar  vigor  of  concoction;"  and 
Harvey  continues:  "Why  Indeed,  should  we  not  as  well  believe  that  the  chyle 
(digested  contents  of  the  intestines)  enters  the  mouth  of  the  mesenteric  veins 
and  in  this  way  becomes  immediately  mingled  with  the  blood,  where  it  might 
receive  digestion  and  perfection.  *  *  *  ^^(j  ^j^^t  the  thing  is  so  in  fact,  I 
find  an  argument  in  the  distribution  of  innumerable  arteries  and  veins  to  the 
intestines,  more  than  to  any  other  part  of  the  body,  in  the  same  way  as  the 
uterus  abounds  in  blood  vessels  during  the  period  of  pregnancy." 

Sir  William  Osier  (Harveian  oration,  1906)  refers  to  this  inci- 
dent in  Harvey's  career:  "How  eminent  so  ever  a  man  may  become 
in  science,  he  is  very  apt  to  carry  with  him  errors  which  were  in 
vogue  when  he  was  young — errors  that  darken  his  understanding, 
and  m.ake  him  incapable  of  accepting  even  the  most  obvious  truths. 
It  is  a  great  consolation  to  know  that  Harvey  came  within  the  range 
of  this  law — in  the  matter  of  the  lymphatic  system;  it  is  the  most 
human  touch  in  his  career." 

The  lacteals  were  demonstrated  in  man  in  1628,  the  subject  be- 
ing an  executed  criminal  examined  shortly  after  execution.  Twenty- 
one  years  after  Aselli's  death  the  thoracic  duct  was  discovered  by 
Johannes  Pecquet,  of  Dieppe,  France.  He  not  only  accurately  de- 
scribed these  lymphatic  structures,  but  showed  that  Aselli's  lacteals 
poured  their  contents  into  what  he  called  the  receptaculum  chyli,  but 
that  the  thoracic  duct — a  continuation  of  the  receptacle — poured  its 
contents  into  the  venous  system  at  the  junction  of  the  jugular  and 
subclavian  veins.  Pecquet  was  twenty-five  years  old  when  he  made 
this  discovery,  which  he  himself  described  as  the  gift  of  fortune 
sporting  with  the  ignorant.  Munus  est  fortunae  cum  inscio  ludentis. 
Pecquet,  however,  did  not  follow  up  this  solitary  triumph.  His  ap- 
petite for  alcoholic  beverages  got  the  better  of  him  and  eventually 
caused  his  death. 

Harvey's  work  on  the  circulation  appeared  between  the  discov- 
ery of  Aselli  and  that  of  Pecquet  and  so  profoundly  had  it  influenced 
the  medical  thought  of  the  time,  that  the  discovery  of  the  thoracic 
duct  and  its  function  was  accepted  without  question. 


C  H  A  P  T  E  R  II. 


PHYSIOLOGY  OF  DIGESTION  IN  THE  SEVENTEENTH 
AND  EIGHTEENTH  CENTURIES 

The  circulation  of  the  blood  was  worked  out  and  proclaimed  to 
the  world  by  one  man,  and  his  work  was  so  complete  that  it  has  not 
been  rendered  obsolete  by  subsequent  knowledge.  The  history  of  the 
physiology  of  digestion  has  been  of  gradual  growth  so  that  no  one 
man  can  claim  credit  for  our  present  knowledge.  Before  the  develop- 
ment of  chemistry,  any  marked  progress  in  the  physiology  of  alimen- 
tation would  not  have  been  possible;  the  early  workers  in  this  par- 
ticular field  were  chemists  rather  than  physiologists.  The  history 
of  physiology  during  the  seventeenth  and  eighteenth  centuries  in- 
volves the  lives  and  work  of  numerous  investigators,  each  accomplish- 
ing all  that  was  possible  considering  the  advancement  of  the  general 
scientific  knowledge  of  the  time. 

Two  names  of  the  latter  part  of  the  seventeenth  and  early  part 
of  the  eighteenth  century  are  prominent  as  exerting  important  infi- 
ence  in  the  way  of  solution  of  the  chemical  problems  of  physiology. 
These  were  George  Ernest  Stahl  and  Hermann  Boerhaave.    Stahl  was 
born  at  Anspach  in  1660 ;  he  studied  at  Jena,  and  after  graduating  be- 
came court  physician  at  Weimer,  and  in  1694  professor  of  medicine  at 
Halle.    He  died  in  1734  in  Berlin,  where  he  moved  in  1716  on  his  ap- 
pointment as  physician  to  the  King  of  Prussia.     Stahl  was  an  ac- 
complished chemist  of  his  day.    His  views  on  gastric  digestion  may  be 
summed  up  in  the  following  sentence  from  his  work:  "Some  people 
suppose  that  gastric  digestion  results  from  the  action  of  particular 
and  specific  ferments,  and  indeed  go  so  far  as  to  regard  the  stomach 
as  not  only  the  seat  but  also  the  origin  of  a  particular  ferment, 
whereas  in  the  whole  construction  of  the  stomach  nothing  particular 
is  observed  which  would  render  the  elaboration  of  such  a  special 
agent  likely."    He  was  a  firm  believer  in  the  psyche  of  Aristotle  and 
introduced  a  principle  which  he  termed  anima.    He  was  wholly  out  of 
sympathy    with    those    who    tried    to    explain    the    physical    and 
psychical  phenomena  of  life  and  mind  on  chemical  and  mechanical 
principles.    He  could  not  think  of  himself  as  a  chemical  retort  subject 
to  ferments.    The  soul  was  to  him  the  living  force  of  the  body;  "It 
was  susceptible  of  being  played  upon  by  a  thousand  different  influ- 
ences, such  as  joy,  sorrow  and  grief,  love  and  friendship,  the  beauti- 
ful, the  true,  the  reverent,  the  sublime.  *  *  *  Can  these  things  be  the 
product  of  chemical  acids  and  alkalies  and  the  mechanical  devices  of 
the  mason  and  builder  ?"    Sir  Michael  Foster  sums  up  the  teaching  of 
Stahl  thus :    "Learn  as  much  as  you  can  of  chemical  and  physical  pro- 
cesses, and  in  so  far  as  the  phenomena  of  the  living  body  exactly  re- 
semble chemical  and  physical  events  appearing  in  non-living  bodies, 
you  may  explain  them  by  chemical  and  physical  laws.     But  do  not 
conclude  that  that  which  you  see  taking  place  in  a  non-living  body 
will  take  place  in  a  living  body,  for  the  chemical  and  physical  phe- 
nomena of  the  latter  are  modified  by  the  soul.    The  events  of  the  bod^ 
may  be  rough  hewn  by  chemical  and  physical  forces,  but  the  soul  will 


PATHFINDERS  OF  PHYSIOLOGY  17 

shape  them  to  its  own  end  and  will  do  that  by  its  own  instrument,  mo- 
tion." Stahl,  it  will  be  seen,  belonged  to  the  "vitalists,"  which  par- 
ticular type  of  physiologist  has  only  within  recent  years  become  ex- 
tinct. His  fundamental  position  was,  between  living  and  non-living 
things  there  is  a  great  gulf  fixed.  Living  things  so  long  as  they  are 
alive  are  actuated  by  the  sensitive  soul;  non-living  things  are  not. 
The  rational  soul  of  man  governed  his  whole  body.  The  healing 
power  of  nature,  vis  medicatrix  naturae,  has  been  recognized  from 
the  most  ancient  to  the  present  time.  Stahl's  system  was  founded 
upon  the  supposition  that  the  vis  naturae  existed  entirely  in  the  ra- 
tional soul.  In  consequence  of  Stahl's  doctrine,  he  and  his  followers 
proposed  the  art  of  curing  by  expectation,  medicina  expectans,  which 
practice  led  to  the  prescribing  of  inert  remedies,  placebos. 

Payer  and  Brunner — In  the  catalogue  of  workers  in  physiology  of 
the  seveneteenth  century  are  the  names  of  Jean  Conrad  Peyer  and 
Brunner.  Peyer  was  bom  in  Switzerland  in  1653.  He  studied  at 
Basel  and  Paris  and  returned  to  his  native  town,  Schaffhausen,  to 
practise,  where  he  died  in  1712.  In  1677  he  published  a  brochure  in 
which  he  described  certain  new  glands  scattered  over  the  intestine; 
these  glands  are  familiar  to  every  student  of  physiology  or  histology 
as  "Peyer's  patches."  He  was  the  first  to  give  a  full  description  of 
these  glnds  are  familiar  to  every  student  of  physiology  or  histology 
lower  part  of  the  small  intestine  and  in  the  ileum,  making  a  distinction 
between  the  single  or  solitary  and  the  patches  of  agminated  glands. 
His  discovery  harmonized  with  that  of  Brunner  a  few  years  later. 

Brunner  was  born  at  Dieffenhausen  in  1653.  He  studied  at 
Strassburg  and  was  eventually  called  to  the  chair  of  medicine  at  Hei- 
delberg, shortly  after  entering  upon  his  position  he  published  his 
Dissertatio  Inauguralis  de  Glandulis  Duodeni,  in  which  he  describes 
the  glands  which  have  since  borne  his  name,  Brunner's  glands.  He 
attributed  to  these  glands  a  function  similar  to  the  pancreas  and 
spoke  of  them  as  a  "pancreas  secondarium."  Brunner  had  made  num- 
erous experiments  by  removing  the  pancreas  from  dogs.  He  con- 
cluded that  the  animals  thus  operated  suffered  in  no  wise  from  ill 
health,  consequently  the  digestive  powers  of  pancreatic  juice  were 
practically  nothing.  These  gropings  of  the  seventeenth  century  are 
curiously  interesting  viewed  in  the  light  of  the  twentieth.  The  work 
of  Peyer  and  Brunner  served  to  deprive  of  its  glory  that  of  Sylvius 
and  DeGraaf,  who  had  attributed  important  digestive  powers  to  the 
pancreatic  juice.  The  attention  of  physiologists  was  again  centered 
on  the  older  view  that  the  stomach  was  the  chief  seat  of  digestion. 

Mechanical  and  Chemical  Views  of  Digestion. — Two  views  con- 
cerning gastric  digestion  contended  for  first  place.  One,  which  may 
be  designated  the  mechanical,  was  espoused  by  Borelli,  who  was  the 
founder  of  the  so-called  latromathematical  school,  which  professed  to 
be  able  to  reduce  all  the  motions  and  activities  of  nature  to  mathe- 
matical formulae.  BorelH's  studies  were  made  on  the  stomachs  or 
gizzards  of  lairds.  He  pointed  out  the  great  grinding  or  pressing  force 
effected  by  the  muscular  coats  of  the  stomach.  He  compares  the  ac- 
tion of  the  fleshy  stomach  to  that  of  the  teeth,  and  continues :  "We 
have  already  shown  that  the  absolute  force  of  the  muscles  which  close 
the  human  jaw  represents  a  power  greater  than  that  of  a  weight  of 
1,350  pounds ;  therefore,  the  force  gf  the  turkey's  stomach  is  not  less 


18  BOERHAAVE 

than  the  power  of  1,350  pounds."  This  estimate  of  the  power  of  the 
human  muscles  of  mastication,  is  rather  high.  Canon  in  his  recent 
work  places  the  pressure  which  the  molars  are  capable  of  exerting  at 
270  pounds.  Borelli  admits,  however,  that  certain  animals  "consume 
flesh  and  bone  by  means  of  a  certain  very  potent  ferment,  much  in 
the  same  way  as  corrosive  liquids  dissolve  metals."  The  iatro-phys- 
ical  school  eventually  went  farther  than  Borelli  and  denied  that  chem- 
ical action  has  anything  whatsoever  to  do  with  digestion,  and  con- 
tended that  digestion  was  mere  trituration  of  the  food  in  the  stomach 
to  a  creamy  substance  known  as  chyle.  Bellini,  a  pupil  of  Borelli? 
went  farther  in  the  beginning  of  the  eighteenth  century  and  endeav- 
ored to  explain  many  functions  of  the  human  body  from  mathemati- 
cal data.  Keill,  a  member  of  this  cult,  calculated  from  data  purely 
imaginary  the  power  of  each  organ.  According  to  him  the  stomach 
had  a  force  of  compression  so  great  that  to  overcome  its  own  resist- 
ance must  have  meant  its  own  destruction.  One  iatro-physicist  esti- 
mated the  force  of  the  heart  as  equal  to  180,000  pounds;  another 
placed  it  at  eight  ounces.  Their  calculations  were  clothed  in  the  im- 
posing nomenclature  of  the  exact  sciences.  This  doctrine  is  said  to 
have  extended  to  all  the  universities  and  medical  institutions  of 
Europe. 

The  iatro-chemical  school,  or  "chemikers"  as  they  were  dubbed 
by  Guy  Patin,  a  French  physician  and  wit  of  the  time,  sought  a  solu- 
tion of  all  the  phenomena  of  the  human  body  in  their  flasks  and  re- 
torts. They  maintained  that  the  change  in  the  stomach  was  chiefly 
if  not  wholly  a  chemical,  resulting  from  the  process  of  fermentation. 
It  was  recognized  even  at  this  time  that  the  membrane  of  the  stomach 
was  glandular  in  structure,  and  yet  little  importance  was  attached  to 
the  secretion  of  such  membraneous  surface. 

In  1614  was  born  Francois  de  le  Boe  or  Dubois,  better  known  by 
his  Latin  name,  Sylvius.  He  is  not  to  be  confused  with  Jacobus  Syl- 
vius, the  Parisian  anatomist,  teacher  of  Vesalius,  who  lived  in  the 
sixteenth  century.  The  latter  Sylvius  studied  at  Sedan  and  at  Basel, 
where  in  1637  he  took  his  degree.  He  became  professor  of  medicine 
at  Leyden,  where  he  exerted  a  powerful  influence  until  his  death  in 
1672.  Sylvius,  though  distinguished  as  a  physician  and  physiologist, 
was  essentially  a  chemist.  Through  his  efforts  the  curators  of  the 
Universtiy  of  Leyden  built  for  him  a  "Laboratorium"  which,  so  far 
as  we  know,  was  the  first  university  chemical  laboratory.  He  devoted 
a  large  part  of  his  time  to  a  study  of  salts,  which  he  learned  to  rec- 
ognize as  resulting  from  the  union  of  acids  with  bases.  Sylvius  looked 
upon  the  phenomena  of  Hfe  from  a  chemical  point  of  view.  He  was 
well  versed  in  that  part  of  physiology  derived  by  deductions  from  an- 
atomy and  by  experiments  on  animals.  His  opinions  on  the  circula- 
tion and  respiration  were  orthodox  from  our  modem  viewpoint.  Har- 
vey's teachings  entered  largely  into  his  thoughts  and  it  was  chiefly 
through  his  advocacy  that  the  doctrine  of  the  great  discoverer  of  the 
circulation  of  the  blood  became  established  in  Holland.  The  contribu- 
tions which  Sylvius  made  to  science  were  essentially  chemical. 

Boerhaave — Herman  Boerhaave,  aready  mentioned  as  a  contrib- 
utor to  the  chemical  knowledge  of  alimentation,  was  born  in  1668, 
near  Leyden,  where  he  was  educated.  His  early  years  were  largely 
devoted  to  the  classical  and  oriental  studies.  He  became  Ph.  D.  in  1690, 


PATHFINDERS  OF  PHYSIOLOGY  19 

having  obtained  the  degree  on  a  thesis,  the  subject  of  which  was  "The 
Distinction  Between  Body  and  Mind."    An  illness  in  the  shape  of  an 
obstinate  ulcer  of  the  leg  turned  his  attention  to  medicine,  which  he 
studied  along  with  the  ancillary  studies,  chemistry  and  botany.     He 
was  graduated  M.  D.  in  1693,  and  eventually  gave  up  the  idea  of  the- 
ology for  medicine.    In  1701  he  was  appointed  to  the  chair  of  medicine 
in  the  University  of  Leyden.    His  great  ability  as  teacher  caused  stu- 
dents to  flock  to  his  lectures.    His  worth  was  quickly  recognized  by 
the  authorities  of  the  university  who  increased  his  emolument  and 
endeavored  to  make  his  position  attractive  to  prevent  him  from  going 
elsewhere.    Sir  Michael  Foster  says  of  him:  "Much  sought  after  as 
a  physician,  acute  at  the  bedside,  brilliant  as  an  expositor  in  the  pro- 
fessorial chair,  he  was  also  a  great  teacher  in  the  sense  that  in  his 
daily  intercourse  with  his  pupils  he  was  always  ready  to  lay  his  mind 
open  before  them  and  to  let  them   share   his  experience  and  his 
thoughts.    Russell  pays  the  following  tribute  to  Boerhaave's  genius : 
"Boerhaave    was    easily    the    most    remarkable    physician    of    his 
age,  a  man  who,  when  we  contemplate  his  genius,  his  condition,  the 
singular  variety  of  his  talent,  his  unfeigned  piety,  his  spotless  char- 
acter and  the  impress  he  left  not  only  on  contemporary  practice,  but 
on  that  of  succeeding  generations,  stands  forth  as  one  of  the  brightest 
names  on  the  pages  of  medical  history,  and  may  be  granted  as  an  ex- 
ample not  only  to  physicians  but  to  mankind."     Boerhaave  was  a 
scholar  and  scientific  thinker,  too  broad  to  be  the  slave  of  one  idea. 
He  was  eclectic  in  the  true  sense  of  the  term,  though  he  never  allied 
himself  with  the  medical  sect  which  goes  by  that  name.     He  had  a 
mind  open  to  truth  wherever  it  might  be  sought.     He  made  use 
of  anatomy,  physics  and  chemistry,  but  never  allowed  one  to  exclude 
the  other.    He  made  each  subservient  to  the  elucidation  of  physiology. 
Boerhaave  was  not  an  extreme  advocate  of  either  mechanical  or 
the  chemical  fermentative  school;  he  recognized  that  digestion  is  in 
part  a  solution  of  some  of  the  constituents  of  food  by  means  of 
various  juices,  which  he,  however,  regarded  not  of  the  nature  of 
fermentation.    He  denied,  however,  the  acidity  of  the  gastric  juice. 
Colored  vegetable  juices  were  at  the  time  coming  to  be  used  as  we 
now  use  litmus  paper,  in  reaction  tests.     Boerhaave  regarded  the 
solution  by  means  of  juices  only  as  part  of  the  digestive  process ;  the 
remaining  process  he  held  consisted  of  trituration  in  the  stomach,  by 
which  process  the  nutritive  parts  of  food  were  expressed.    His  views 
were  dominant  the  early  part  of  the  eighteenth  century. 

An  Epochal  Year,  1757 — The  years  1757  was  the  dividing  line  be- 
tween modern  physiology  and  all  that  had  gone  before.  It  was  the 
date  of  the  publication  of  the  first  volume  of  Haller's  Elementa  Phys- 
iologia,  the  eighth  volume  of  which  appeared  in  1765.  Albrecht  von 
Haller  was  born  at  Berne,  Switzerland,  in  1708.  The  story  is  told  of 
his  early  precosity,  when  at  the  age  of  four  he  is  said  to  have  ex- 
pounded the  Bible  to  his  father's  servants.  Before  he  was  ten,  he 
wrote  in  Latin  verse  a  satire  on  his  tutor,  Haller's  attention  had  been 
directed  to  medicine  after  his  father's  death  in  1721,  while  residing  in 
the  house  of  a  physician  in  Biel,  and  in  his  sixteenth  year  he  entered 
the  University  of  Tubingen.  Dissatisfied  with  his  progress  there,  he 
went  to  Leyden  ,where  Boerhaave  was  at  the  height  of  his  fame.  He 
graduated  in  1727,  and  turned  his  attention  to  botany,  publishing  a 


20  HALLBR 

great  work  on  the  flora  of  Switzerland.  He  returned  to  Berne  and  be- 
gan the  practice  of  medicine  in  1729.  In  1736  he  was  appointed  pro- 
fessor of  medicine,  anatomy  and  botany  in  the  newly  founded  univer- 
sity of  Gottingen,  a  position  which  he  had  held  for  17  years.  During 
tljjs  time  he  carried  on  original  inve^tigat<on  in  botany  and  physiology. 
His  researches  on  the  formation  of  bone,  the  mechanics  of  respiration, 
and  the  development  of  the  embryo  are  of  the  highest  importance.  Re- 
garding Haller  as  an  expositor  in  physiology,  Foster  writes :  "When 
we  turn  from  the  preceding  writers  on  physiology  and  open  the  pages 
of  Haller's  Elementa,  we  feel  that  we  pass  into  modem  times.  Save 
for  the  strangeness  of  most  of  the  nomenclature,  and  for  no  small 
differences  in  all  that  relates  to  the  chemical  changes  of  the  body,  we 
seem  to  be  reading  a  modem  text-book  of  the  most  exhaustive  kind." 
His  chief  service,  however,  was  the  careful  arranging  and  digesting 
of  the  theories  and  facts  of  physiology  up  to  this  time.  From  his  time 
physiology  became  an  independent  branch  of  science,  to  be  pursued 
for  itself  rather  than  as  an  adjunct  to  medicine.  Regarding  Haller's 
method  of  exposition,  the  same  writer  goes  on  to  say  that  "In  dealing 
with  each  subdivision  of  physiology,  Haller  carefully  describes  the 
anatomical  basis,  including  the  data  of  minute  structure,  physical 
properties  and  chemical  composition,  so  far  as  these  were  then  known. 
He  then  states  the  observations  that  have  been  made,  and  in 
respect  to  each  question,  as  it  arises,  explains  the  several  views 
which  have  been  put  forward,  giving  minute  and  full  references  to 
all  the  authors  quoted,  and  he  finally  delivers  a  reasoned  critical  judg- 
ment expounding  the  conclusions  which  may  be  arrived  at,  but  not 
omitting  to  state  plainly  when  necessary  the  limitations  which  the 
lack  of  adequate  evidence  places  on  forming  a  decided  judgment.  He 
carefully  recounts  and  as  carefully  criticizes  all  the  knowledge  that 
can  be  gleaned  about  any  question.  If  he  feels  unable  to  come  to  a 
decided  conclusion  he  candidly  says  so." 

But  we  are  most  concerned  at  present  with  what  Haller  has  to 
say  on  digestion.  He  considered  saliva  neutral  in  reaction  and  pos- 
sessing no  digestive  properties  further  than  the  softening  of  food  as 
an  aid  to  deglutition.  He  recognized  the  importance  of  the  glandular 
coat  of  the  stomach,  which  glands  he  concluded  furnished  mucous 
only,  the  true  gastric  juice  being  derived  from  the  arteries.  He  also 
concluded  that  pure  gastric  juice  was  neither  acid  nor  alkaline  and 
refused  to  regard  it  as  some  of  his  predecessors  had  done,  as  a  fer- 
ment. The  acidity  he  considered  a  token  of  the  degeneration  of  the 
digested  food.  Trituration  he  regards  as  a  useful  aid  to  digestion,  es- 
pecially where  hard  grains  form  part  of  the  food  as  in  birds;  but  it 
was  only  an  aid. 

Bile,  he  claimed,  was  not  a  mere  excrement;  it  was  secreted  by 
the  liver  and  stored  for  a  time  in  the  gall  bladder,  where  it  underwent 
slight  change.  Bile  is  a  viscid  fluid,  bitter  but  neither  acid  nor  alka- 
line. It  has  the  power  of  dissolving  fats,  and  so  acts  on  a  mixture 
of  oil  and  water  as  to  form  an  emulsion.  Haller  considered  the  im- 
portance of  the  pancreas  due  to  the  fact  that  its  ducts  opened  into  the 
intestine  in  common  with  the  bile  duct;  that  its  fluid  softened  and 
diluted  the  bile,  thus  enabling  it  to  mix  more  satisfactory  with  the 
food.  He  concluded  by  prophesying  that  there  may  be  other  func- 
tions of  pancreatic  juice  not  well  known  to  the  physiologists  of  his 
day. 


PATHFINDERS  OF  PHYSIOLOGY  21 

Reaumur  and  His  Methods.  Rene  Antoine  Ferchault  de  Reaumur, 
a  Frenchman  bom  in  1683,  and  described  as  one  of  the  most  notable 
men  of  science  of  the  eighteenth  century,  is,  in  chronological  se- 
quence next  most  important  contributor  to  the  physiology  of  the 
alimentary  tract.  His  name  is  already  famihar  to  most  of  us  as  the 
inventor  of  the  Reaumur  thermometer.  His  studies  on  the  gastric 
juice  at  this  time  are  all-important,  inasmuch  as  his  methods  are 
unique.  Reaumur  had  in  his  possession  a  kite  and  took  advantage  of 
the  habit  of  the  bird  of  ejecting  from  its  stomach  things  swallowed 
which  it  could  not  digest.  The  kite  was  fed  pieces  of  meat  secured 
in  metal  tubes.  It  was  found  that  meat  when  ejected  had  no  odor 
of  putrification.  Experiments  were  made  with  small  pieces  of  bone, 
which  were  completely  dissolved  when  ejected  and  swallowed  by  the 
kite  several  times.  On  vegetable  grains  and  flour,  the  fluid  of  the 
kite's  stomach  had  apparently  little  effect.  The  tubes  were  filled  with 
small  pieces  of  sponge,  which,  when  ejected,  were  squeezed  out,  thus 
enabling  the  investigator  to  procure  pure  gastric  juice  and  to  study 
it  in  vitro.  He  proved  that  digestion  was  not  putrifaction  but  some- 
thing really  opposed  to  that  process.  While  Reaumur's  experiments 
left  much  to  be  ascertained  about  gastric  digestion,  he  at  least  favored 
the  solvent  power  of  the  succus  gatricus,  by  the  employment  of  a 
wholly  new  method. 

Experiments  with  Gastric  Juice.  We  must  look  to  Italy  for  the 
next  contributor  to  our  knowledge  of  digestion.  Parenthetically,  it 
is  of  interest  to  note  that  the  idea  of  specializing,  if  it  had  taken  root 
at  all  at  this  early  time,  was  not  markedly  apparent.  The  worker  in 
the  physiology  of  digestion  was  equally  prominent  in  almost  every 
other  department  of  physiological  research.  Lazzaro  Spallanzani 
(1729-1799)  was  one  of  the  most  eminent  men  of  his  time.  Educated 
for  the  church,  he  was  usually  known  as  Abbe  Spallanzani.  His  life 
was  devoted  to  experiments,  researches  and  teaching.  He  was  pro- 
fessor at  Bologna,  and  afterwards  at  Pavia.  We  find  him  first  ex- 
perimenting with  germ  life,  with  results  that  disprove  the  doctrine 
of  spontaneous  generation.  His  researches  in  other  fields  showed  that 
he  had  conceived  the  truly  scientific  method. 

Spallanzani  took  up  Reaumur's  methods  and  most  of  his  results 
were  achieved  by  them.  Aided  by  improvements  in  chemistry,  he 
was  able  to  make  marked  advance  over  his  predecessors.  His  ex- 
periments were  made  on  all  kinds  of  animals,  fishes,  frogs,  serpents, 
birds,  sheep  oxen,  horses,  cats  and  dogs,  and  lastly  upon  himself.  Be- 
sides hollow  tubes,  he  used  hollow  spheres,  freely  perforated,  into 
which  were  placed  meat  and  bread,  bone  or  grains  of  wheat,  and  the 
results  of  digestion  were  studied  when  these  were  ejected  or  procured 
by  opening  the  animal's  stomach.  He  also  attached  pieces  of  meat  to 
threads,  which  he  would  draw  from  the  animal's  stomach  at  fixed  in- 
tervals. He  experimented  upon  himself  by  swallowing  linen  bags  con- 
taining bread,  meat  and  similar  articles,  examining  the  contents  after 
they  had  been  voided  per  anum.  He  procured  gastric  juice  from 
himself  by  producing  vomiting  on  an  empty  stomach.  He  repeatedly 
tested  the  action  of  gastric  juice  in  vitro,  keeping  the  tubes  a  uniform 
temperature  by  retaining  them  in  his  crm  pit,  using  the  same  food 
covered  by  water  as  a  control.  He  found  that  gastric  juice  acted 
more  readily  upon  finely  divided  parts  of  food  such  as  crushed  grain 


22  SPALLANZANI 

or  bone  which  proved  trituration  only  a  preparation  for  solution,  and 
that  it  was  no  further  a  part  of  the  digestive  process. 

He  found  that  the  gastric  juice  dissolved  the  food  of  animals  into 
a  pultaceous  mass  or  chyme.  He  observed  that  heat  favored  solution 
and  that  in  warm-blooded  animals  certain  high  temperature  was 
necessary  for  the  chymification  of  foods.  In  Spallanzani's  time  putri- 
faction  was  considered  a  form  of  fermentation.  "There  are  three 
kinds  of  fermentation,  the  vinous,  the  acetous,  and  the  putrid."  The 
action  of  gastric  juice  was  not  putrid;  in  fact,  it  tended  to  arrest  put- 
refaction. Spallanzani  was  inclined  to  believe  that  the  action  of  the 
gastric  juice  was  neither  vinous  nor  acetous.  Regarding  the  reaction 
of  gastric  juice  his  conclusion  was  that  it  was  neutral.  He  believed 
that  the  acidity  was  due  to  an  abnormality  of  the  stomach  contents, 
inasmuch  as  the  regurgitation  of  sour  material  from  the  stomach  oc- 
curred only  when  something  had  gone  wrong.  Spallanzani's  failure  to 
recognize  the  acidity  of  the  gastric  juice  limited  his  further  investiga- 
tions. He  could  only  conclude  that  the  action  of  the  gastric  juice  was 
not  fermentation,  as  fermentation  was  understood  at  the  time. 

It  is  interesting  to  note  that  the  results  of  Reaumur  and  Spallan- 
zani were  confirmed  by  Stevens  of  Edinburgh,  who  likewise  employed 
Reaumur's  methods  of  investigation.  Stevens  experimented  on  a 
"man  of  weak  understanding  who  gained  a  miserable  livelihood  by 
swallowing  stones  for  the  amusement  of  the  common  people."  The 
man  was  made  to  swallow  perforated  silver  spheres  containing  animal 
and  vegetable  food,  raw  and  cooked,  which  were  examined  when  void- 
ed some  48  hours  later.  Similar  experiments  were  made  on  dogs,  the 
contents  of  the  hollow  spheres  examined  after  opening  the  animal's 
stomach.  Stevens  concluded  that  digestion  is  not  the  effect  of  heat, 
trituration,  putrification  or  fermentation  alone,  but  of  a  powerful  sol- 
vent secreted  by  the  glandular  coat  of  the  stomach. 

Summary:  Summing  up  the  progress  made  in  the  physiology  of 
digestion  during  the  seventeenth  and  eighteenth  centuries,  probably 
no  one  is  more  entitled  to  an  audience  than  Sir  Michael  Foster;  "Dur- 
ing the  two  centuries  the  seventeenth  and  the  eighteenth,  physiologi- 
cal inquiries,  swayed  now  in  one  direction  by  views  of  chemical  fer- 
mentation or  effervescence,  now  in  another  direction  by  views  of 
mechanical  trituration,  had  come  in  the  end  to  the  conclusion  that 
digestion  was  in  the  main  a  process  of  solution  of  a  peculiar  charac- 
ter, begun  and  chiefly  carried  out  in  the  stomach,  though  assisted  by 
minor  subsequent  changes  taking  place  along  the  intestines.  They 
who  were  under  the  influence  of  vitalistic  doctrines,  and  these  were 
perhaps  the  more  numerous,  held  the  change  to  be  the  commencement 
of,  to  be  the  first  step  in  the  conversion  of  food  into  living  flesh  and 
blood,  and  spoke  of  it  as  a  change  differing  from  ordinary  chemical 
change,  without  being  able  to  define  the  exact  characters.  It  was  left 
to  the  nineteenth  century  to  throw  new  light  on  the  nature  of  the 
gastric  changes  and  at  the  same  time  to  show  that  what  took  place 
in  the  stomach  was  not  the  whole  digestion,  but  only  the  first  of  a 
series  of  profound  changes  taking  place  along  nearly  the  whole  length 
of  the  alimentary  canal." 


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CHAPTER  III. 


PHYSIOLOGY  OF  DIGESTION— WILLIAM  BEAUMONT 

We  have  traced  the  development  of  the  physiology  of  alimenta- 
tion from  its  crude  beginnings,  when  debate  waged  as  to  whether 
digestion  consisted  of  mechanical  trituration  or  whether  it  consisted 
wholly  of  a  fermentative  process,  to  the  time  when  some  real  light 
began  to  be  shed  upon  the  subject  by  experiment  with  the  gastric 
secretion  itself.  No  contribution  to  the  subject  of  gastric  digestion 
has  been  of  such  moment  as  the  work  of  William  Beaumont  on  the 
gastric  secretion  of  the  French  Canadian,  Alexis  St,  Martin.  The  story 
of  Beaumont's  life  and  the  circumstances  surrounding  his  work  con- 
stitute one  of  the  most  fascinating  chapters  in  the  history  of  Ameri- 
can Medicine. 

As  physicians,  we  have  become  familiar  with  the  names  of  Beau- 
mont and  St.  Martin  early  in  our  student  career.  They  have  become 
inseparably  associated  with  the  study  of  gastric  juice  and  its  func- 
tions. Standard  works  on  physiology  introduce  the  chapter  on  diges- 
tion with  such  sentences  as:  "Gastric  fistulae  have  been  made  in 
human  beings,  either  by  accidental  injury  or  by  surgical  operation. 
The  most  celebrated  case  is  that  of  Alexis  St.  Martin,  a  young  Can- 
adian who  received  a  musket  wound  in  the  abdomen  in  1822.  Obser- 
vations made  upon  him  by  Dr.  Beaumont  formed  the  starting  point  of 
our  correct  knowledge  of  the  physiology  of  the  stomach  and  its  secre- 
tions."* "The  first  fistula  of  a  digestive  gland  to  be  the  subject  of  a 
thoroughly  scientific  investigation  was  one  resulting  from  a  gun  shot 
wound  in  the  stomach  of  a  Canadian  hunter.  As  the  consequence  of 
his  accident,  the  hunter  had  all  the  rest  of  his  life  a  stomach  fistula 
opening  at  the  upper  part  of  the  abdomen,  through  which  the  interior 
of  the  stomach  could  be  observed  and  gastric  juice  could  be  obtained. 
Beaumont  collected  a  large  number  of  important  facts  (1825-1833) 
concerning  the  digestive  process  of  the  stomach  and  concerning  the 
movements  of  that  organ."  "Beaumont's  studj^  of  St.  Martin's  stom- 
ach showed  that  in  acute  catarrh  the  mucous  membrane  is  reddened 
and  swollen,  less  gastric  juice  is  secreted,  and  mucous  covers  the  sur- 
face." Instances  might  be  quoted  almost  ad  infinitum  of  references 
in  medical  literature  to  Beaumont's  classic  study  of  gastric  digestion. 

Beaumont ;  His  Early  Life :  William  Beaumont,  the  third  child  of 
Samuel  Beaumont,  who  had  seen  active  service  during  Revolution  days 
prior  to  the  Declaration  of  Independence,  was  born  November  21st, 
1785.  There  was  nothing  unusual  in  his  childhood  and  youth.  As  he 
grew  to  manhood  his  sympathies  and  political  leanings  were  in  accord 
with  those  of  his  father,  who  was  a  staunch  Democrat  and  patriot. 
While  no  church  record  assures  us  that  he  was  of  the  faith  of  his 
parents,  Congregationalist,  his  biographer  asserts  that  when  the  roll 
of  the  drum  announced  the  approaching  hour  of  worship  he  was 
among  those  who  slowly  wended  their  way  over  the  hills  on  foot  or  on 

♦Haliburton's  Handbook  of  Physiology;     Tigerstedt's    Physiology;     Osier's 
Practice. 


24  BEAUMONT 

horseback  to  the  old  meeting  house.  Beaumont  was  blessed  with  such 
rigorous  parental  discipline  in  youth  that  he  explained  his  lapses  in 
church  attendance  in  after  life  by  the  statement  that  during  his 
youth  he  had  made  up  for  a  lifetime  of  church  attendance.  Further 
than  that  he  was  a  courageous  and  fearless  boy,  little  is  known  of  his 
early  life.  It  is  said  that  he  developed  deafness,  which  became  more 
marked  as  he  grew  older,  from  standing  near  a  cannon  which  was 
being  fired,  simply  to  outwit  playmates  of  his  own  age. 

The  beginning  of  last  century  found  Beaumont  a  boy  of  fifteen 
years.  It  was  twenty-four  years  since  the  first  birthday  of  the  Ameri- 
can Nation.  Beaumont's  youth  was  contemporaneous  with  one  of  the 
most  stirring  epochs  in  world  history.  The  United  States  was  begin- 
ning to  assume  an  important  place  among  the  nations  of  the  world. 
Beaumont  left  home  during  the  winter  of  1806-7  with,  we  are  told,  an 
outfit  consisting  of  a  horse  and  cutter,  a  barrel  of  cider,  and  a  hundred 
dollars  of  hard  earned  money.  He  traveled  Northward,  reaching  in 
the  spring  of  1807  the  little  village  of  Champlain,  New  York.  He  was 
very  favorably  impressed  with  his  surroundings  and  with  the  people, 
who  were  mostly  farmers,  and  whom  he  characterized  as  "peaceful 
and  industrious  in  general."  Here  he  established  his  "Lares  and 
Penates"  and  followed  the  career  of  schoolmaster.  Coming  from  one 
of  the  best  New  England  schools  his  services  were  much  in  demand. 
While  teaching  school  and  during  the  vacation  he  found  time  to  de- 
vote to  medical  studies.  He  had  supplied  himself  with  books  borrowed 
from  Dr.  Pomeroy  of  Burlington,  Vt.,  which  town  was  on  his  itinerary 
to  Champlain.  Beaumont,  as  many  since  his  day  have  done,  made 
teaching  a  stepping  stone  to  the  profession  of  medicine,  and  an  excel- 
lent experience  it  is  for  the  aspiring  savant.  In  1810  he  was  apprentic- 
ed to  Dr.  Chandler  of  St.  Albans,  Vt.  He  seems  to  have  exhibited  a 
wise  choice  in  the  matter  of  preceptor.  "Living  under  the  same 
roof,"  writes  Dr.  Myer,  describing  the  medical  education  of  the 
times,  "as  was  customary  in  the  days  of  medical  apprenticeship,  the 
preceptor  could  look  after  both  the  mind  and  morals  of  his  pupils. 
The  fledgeling  in  return  for  the  instruction  received  at  the  hands  of 
his  master,  not  only  compensated  him  for  his  trouble,  but  performed 
many  of  the  menial  offices  of  a  servant  about  the  house  and  office. 
It  was  he  who  prepared  the  powders,  mixed  the  concoctions,  made 
pills,  swept  the  office,  kept  the  bottles  clean,  assisted  in  operations 
and  often  through  main  force  supplied  the  place  of  the  anaesthetic  of 
today,  in  the  amputation  of  limbs  and  other  surgical  procedures.  He 
rode  about  with  the  doctor  from  house  to  house,  profiting  by  his  per- 
sonal experience  and  jotting  down  on  the  pages  of  his  note  book  and  on 
the  tablets  of  his  memory  the  words  of  wisdom  that  fell  from  his  mas- 
ter's lips.  *  *  *  He  was  taught  the  symptoms  of  disease,  the 
crude  methods  of  diagnosis,  the  art  of  prescription  writing  and  the 
process  of  cupping  and  bleeding,  considered  so  effective  in  its  day."* 

Medical  books  were  rare  and  expensive,  and  fortunate  was  the 
student  who  had  access  to  them.  Dissections  were  rarely  performed, 
owing  largely  to  the  fact  of  inadequate  means  of  preserving  cadavers. 
Such  were  young  Beaumont's  opportunities. 

Beaumont  spent  the  two  years  of  his  apprenticeship  with  dili- 
gence, studying  the  masters.  He  dissected  whenever  an  opportunity 
♦Life  and  letters  of  Dr.  William  Beaumont,  by  Jesse  S.  Myer. 


BEAUMONT  35 

afforded,  and  never  lost  an  opportunity  to  perform  post-mortems.  A 
perusal  of  his  case  histories  shows  what  a  careful  observer  he  was — 
a  qualification  of  the  first  importance  in  a  physician.  His  diploma  or 
license  to  practice  was  granted  the  second  Tuesday  of  June,  1812,  by 
the  third  Medical  Society  of  the  state  of  Vermont.    It  reads : 

By  the  Third  Medical  Society  of  the  State  of  Vermont  as  by  law  estab- 
lished, William  Beaumont  having  presented  himself  for  examination  on  the  anat- 
omy of  the  human  body,  and  the  theory  and  practice  of  physic  and  surgery,  and 
being  approved  by  our  censors,  the  society  willingly  recommends  him  to  the 
world  as  a  judicious  and  safe  practitioner  in  the  different  avocations  of  the  medi- 
cal profession.  In  testimony  whereof  we  have  hereunto  prefixed  the  signature 
of  our  president  and  the  seal  of  the  society  at  the  Medical  Hall  in  Burlington  on 
the  second  Tuesday  of  June,  A.  D.  1812, 

CASSIUS  F.  POMEROY,  Secretary.  JOHN  POMERY,  President 

Assistant  Army  Surgeon:  In  September  the  same  year  Beau- 
mont joined  the  army  at  Plattsburgh,  as  assistant  surgeon  under  Gen- 
eral Dearborn.  His  old  preceptor  Dr.  Chandler  had  unsuccessfully 
tried  to  dissuade  him  from  the  army  service,  advising  him  to  settle 
down  to  private  practice.  Apparently  there  is  a  destiny  which  shapes 
our  ends.  Had  he  followed  the  advice  of  his  old  master,  he  would  in 
all  probability  have  been  among  the  thousands  of  good  men  who  have 
lived  their  lives  through,  leaving  the  world  a  little  better  than  they 
found  it,  and  passed  into  the  silent  land,  pass  and  leave  no  sign  to  in- 
dicate that  they  have  been.  But  Beaumont  followed  his  ov/n  bent  and 
it  was  while  acting  as  army  surgeon  that  he  made  the  momentous 
discoveries  which  have  placed  him  among  the  epoch-makers  of  medi- 
cal history.  It  is  significant  to  note  that  more  than  one  army  sur- 
geon has  performed  service  of  an  extraordinary  nature  to  medical 
science.  Fom  the  times  when  Machaon  and  Podilirius  rendered  aid  to 
the  Greek  hosts  at  ancient  Troy  to  the  days  of  Ambrose  Pare,  the 
army  surgeon  has  been  identified  with  medical  progress.  A  name 
honored  within  recent  years  in  the  French  service  is  that  of  Laveran, 
who  during  his  tour  of  duty  in  Algeria  did  a  work  in  connection  with 
malaria  which  made  possible  the  work  of  Sir  Ronald  Ross,  of  the  In- 
dian medical  service,  and  his  associates  of  more  recent  times.  The 
spectacle  of  the  Panama  Canal  and  its  construction  v/ere  made  pos- 
sible by  the  United  States  Army  medical  service.  In  the  British  Army 
medical  service  are  such  names  as  Sir  David  Bruce,  whose  investiga- 
tions led  to  the  extermination  of  Malta  fever. 

Beaumont's  Diary:  Beaumont  left  a  diary  which  is  an  interest- 
ing description  by  one  on  the  firing  line,  of  the  stormy  times  of  1812. 
This  graphic  account  of  events  of  the  war  by  an  eye-witness  is 
reproduced  in  Dr.  Meyer's  book.  Beaumont  was  present  August,  1814, 
at  the  battle  of  Plattsburgh,  where  General  Macomb  defeated  the  Brit- 
ish under  General  Provost.  The  Treaty  of  Ghent  ratified  in  February, 
1815,  closed  the  war.  Soon  after  the  close  of  the  war  of  1812  Beau- 
mont tendered  his  resignation  and  in  partnership  with  a  Dr.  Senter 
opened  a  store  in  the  town  of  Plattsburgh,  which  store  contained  "a 
general  assortment  of  drugs,  medicines,  groceries,  dye  woods,  etc.,  of 
the  first  quality  and  choicest  selection  which  they  calculate  to  sell  on 
liberal  terms  for  cash  or  approved  credit."  So  runs  the  advertisement 


26  PATHFINDERS  OF  PHYSIOLOGY 

in  the  local  newspaper.  In  the  footnote  of  the  advertisement  it  is 
stated  that  "Medicines  will  be  put  up  with  accuracy  and  care."  In 
December  of  1816  Beaumont  sold  out  and  afterwards  confined  himself 
entirely  to  the  practice  of  his  profession.  He  was  commissioned  by 
President  Monroe  in  1820  and  re-entered  the  military  service,  when 
he  was  ordered  to  Fort  Mackinac  on  the  Northwestern  frontier.  He 
describes  his  journey  in  detail  in  his  diary.  His  course  lay  along  the 
southern  shore  of  Lake  Erie  to  the  Detroit  river,  where  he  passed 
Fort  Maiden,  near  the  Canadian  town  of  Amherstburg,  opposite  Bois 
Blanc  island.  He  describes  the  fort  at  Detroit  as  a  "regular  work  of 
an  oblong  figure  covering  about  an  acre  of  graceful  slopes."  The 
parapets  are  about  20  ft.  in  height,  built  of  earth  and  sodded,  with 
four  bastions.  The  whole  surrounded  with  palisades,  a  deep  ditch 
and  glacis.  It  stands  immediately  back  of  the  town  and  has  strength 
to  withstand  a  siege.  The  Detroit  postoff  ice,  corner  of  Fort  and  Shelby 
streets,  stands  upon  the  ground  at  one  time  occupied  by  the  above 
mentioned  fortification.  A  bronze  tablet  at  the  south  entrance  of  the 
postoffice  gives  in  brief  the  vicissitudes  of  the  old  fort. 

He  speaks  of  crossing  over  to  Sandwich,  then  a  small  French  vil- 
lage. There  is  no  mention  of  the  route  again  until  he  reaches  Fort 
Michilimackinac,  which  is  described  as  handsomely  situated  on  the 
southeast  side  of  the  island  of  this  name,  on  a  bluff  rising  from  100 
to  200  feet  from  the  water,  almost  perpendicular  in  many  places,  ex- 
tending about  half  way  around  the  island.  The  word  "Michilimacki- 
nac" means  "turtle"  from  the  resemblance  of  Mackinac  island  on  be- 
ing approached. 

The  following  entries  in  his  diary  throw  considerable  light  on  the 
character  of  the  man  himself. 

Sept.  9,  1820.  Commenced  a  diary  of  conduct  on  Dr.  Franklin's  plan,  for  ob- 
taining moral  perfection."  (Benjamin  Franklin  appears  to  have  been  a  favorite 
with  Beaumont,  for  he  elsewhere  quotes  him  at  length.)  "Reading  Shakespeare 
today  I  judged  the  following  extracts  worthy  of  copying;  'Love  all,  trust  few, 
do  wrong  to  none;  be  able  for  thine  enemy  rather  in  power  than  use;  and  keep 
thy  friend  under  thy  life's  key;  be  checked  for  silence,  but  more  taxed  for  speech. 

"10th.  Rose  at  six  o'clock.  Visited  my  patients  in  village  and  discharged 
garrison  duty  before  9  a.  m.  Settled  my  hospital  account,  perused  scriptures 
and  Pope's  Essay  on  Man  till  evening." 

Beaumont's  diary  is  an  interesting  narrative  of  the  times,  written 
by  a  keen  and  practical  observer. 

The  Psychological  Moment:  Late  in  the  spring  of  1822  occurred 
the  event  which  made  the  name  of  William  Beaumont  famous  in  the 
annals  of  medicine.  Indians  and  voyageurs  had  returned  to  Mackinac 
with  the  results  of  the  winter's  hunting.  A  strange  medley  of  hu- 
manity had  gathered  at  the  Amercan  Fur  Company's  trading  post. 
On  the  6th  of  June  a  gun  was  accidentally  discharged,  its  contents 
entering  the  upper  abdomen  of  a  young  voyageur,  leaving  a  cavity 
which  would  have  admitted  a  man's  fist.  According  to  an  eye-wit- 
ness Alexis  St.  Martin,  for  that  was  his  name,  fell,  as  every  one  sup- 
posed, dead.  Dr.  Beaumont,  surgeon  of  the  fort,  was  called,  and  ar- 
rived shortly  after  the  accident.  Shot  and  pieces  of  clothing  were 
extracted  and  the  wound  dressed.    The  surgeon  then  left  with  the  re- 


BEAUMONT  27 

mark  that  the  man  couldn't  live  36  hours.  The  doctor  called  again  in 
the  course  of  two  or  three  hours  and  found  the  patient  better  than  he 
had  anticipated.  The  patient  was  removed  to  the  fort  hospital  where 
he  eventually  recovered,  leaving-,  however,  a  permanent  gastric  fistula. 
Beaumont's  own  account  of  the  accident  is  told  in  the  introduction 
to  his  work  on  "Experiments  and  Observations  of  Gastric  Juice." 

"Alexis  St.  Martin,  who  is  the  subject  of  these  experiments,  was  a  Canadian 
of  French  descent  at  the  above  mentioned  time  (1822)  about  18  years  of  age,  of 
good  constitution,  robust  and  healthy.  He  had  been  engaged  in  the  service  of 
the  American  Fur  Company  as  a  voyager  and  was  accidentally  wounded  by  the 
discharge  of  his  musket  on  the  6th  of  June;  the  charge,  consisting  of  powder  and 
duck-shot,  was  received  in  the  left  side  of  the  youth,  he  being  at  a  distance  of  not 
more  than  one  yard  from  the  muzzle  of  the  gun.  The  contents  entered  posteriorly 
and  in  an  oblique  direction,  forward  and  inward,  literally  blowing  off  integuments 
and  muscles  of  the  size  of  a  man's  hand,  fracturing  and  carrying  away  the  anter- 
ior half  of  the  sixth  rib,  lacerating  the  lower  portion  of  the  left  lung,  the  dia- 
phragm and  perforating  the  stomach.  The  whole  mass  of  materials  forced  from 
the  musket,  together  with  fragments  of  clothing  and  pieces  of  fractured  ribs, 
were  driven  into  the  muscles  and  cavity  of  the  chest.  I  saw  him  in  25  or  30  min- 
utes after  the  accident  occurred,  and  on  examination  found  a  portion  of  the  lung 
as  large  as  a  turkey's  egg  protruding  through  the  external  wound,  lacerated  and 
burned;  and  immediately  below  this  another  protrusion  which,  on  further  ex- 
amination, proved  to  be  a  portion  of  the  stomach  lacerated  through  all  its  coats 
and  pouring  out  the  food  he  had  taken  for  his  breakfast  through  an  orifice  large 
enough  to  admit  the  forefinger." 

Beaumont's  hospital  and  bedside  notes  give  a  complete  history 
of  the  case. 

Being  destitute  and  without  friends  or  relatives,  Alexis  St.  Mar- 
tin became  a  pauper  on  the  town  of  Mackinac.  It  was  at  last  decided 
to  ship  him  to  his  native  town,  Montreal,  nearly  one  thousand  miles 
away.  Beaumont,  however,  rescued  him  from  misery  and  inevitable 
death  by  taking  him  into  his  own  family.  "During  this  time,  says 
his  benefactor,  I  nursed  him,  fed  him,  clothed  him,  lodged  him  and 
furnished  him  with  every  comfort  and  dressed  his  wounds  daily  and 
for  the  most  part  twice  a  day."  It  should  be  realized  that  Beau- 
mont endeavored  to  close  the  wound;  that  when  all  other  means 
failed  he  suggested  incising  the  edges  of  the  wound  and,  "bringing 
them  together  by  sutures,  an  operation  to  which  the  patient  would 
not  submit." 

Not  until  three  years  after  the  accident  did  the  idea  of  perform- 
ing a  number  of  experiments  appear  to  occur  to  the  mind  of  Beau- 
mont. In  1825  he  began  to  realize  the  importance  of  this  case  which 
had  fallen  to  his  care,  when  it  occurred  to  him  what  a  great  service 
to  humanity  might  result  from  this  accident.  About  this  time  Beau- 
mont describes  the  situation  as  follows: 

'He  (St.  Martin)  will  drink  a  quart  of  water  or  eat  a  dish  of  soup  and  then 
by  removing  the  dressings  I  frequently  find  the  stomach  inverted  to  the  size 
and  about  the  shape  of  a  half-blown  rose,  yet  he  complains  of  no  pain,  and  it 
will  return  itself  or  is  easily  reduced  by  gentle  pressure.  When  he  lies  on  the 
opposite  side  I  can  look  directly  into  the  cavity  of  the  stomach  and  almost  see 
the  processes  of  digestion.  I  have  frequently  suspended  flesh,  raw  and  wasted, 
and  other  substances  into  the  perforation  to  ascertain  the  length  of  time   re- 


28  PATHFINDERS  OF  PHYSIOLOGY 

quired  to  digest  each,  and  at  one  time  used  a  tent  of  raw  beef  instead  of  lint 
to  stop  the  orifice,  and  found  that  in  less  than  five  hours  it  was  completely 
digested  off  as  smooth  and  as  even  as  if  it  had  been  cut  with  a  knife." 

Then  his  resolve  to  make  use  of  the  case  as  a  means  of  study- 
ing gastric  digestion  takes  shape  as  follows: 

"This  case  affords  an  excellent  opportunity  for  experimenting  on  the  gastric 
fluid  and  process  of  digestion.  It  would  give  no  pain  nor  cause  the  least  un- 
easiness to  extract  a  gill  of  fluid  every  two  or  three  days  for  it  frequently  flows 
out  spontaneously  in  considerable  quantities.  Various  kinds  of  digestible  sub- 
stances might  be  introduced  into  the  stomach  and  then  easily  examined  during 
the  whole  process  of  digestion.  I  may,  therefore,  be  able  hereafter  to  give  some 
interesting  experiments  on  these  subjects." 

Recognition  of  Michigan  Medical  Society:  The  Medical  Society 
of  the  territory  of  Michigan  was  the  first  body  to  recognize  the  work 
of  William  Beaumont.  The  following  letter  dated  from  Detroit  an- 
nounced his  election  as  an  honorary  member  of  the  Michigan  Territor- 
ial Medical  Society. 

"Dr.  William  Beaumont,  United  States  Army,  Michilimackinac. 

Detroit,  March  3,  1825. 

"Sir: — It  is  with  much  pleasure  that  I  transmit  to  you  as  an  extract  from 
the  minutes  of  the  medical  society  of  this  territory  at  a  meeting  held  at  the  home 
of  Capt.  Woodworth  in  the  City  of  Detroit  on  Monday,  7th  ultimo;  Dr.  VVilliam 
Beaumont,  of  the  United  States  Army,  duly  proposed  by  Dr.  Pitcher  and  unani- 
mously elected  by  ballot  an  honorary  member  of  this  society.' 

"Whereupon   it  was   ordered   that  the  secretary  be   directed   to   Inform  Dr. 
Beaumont  of  his  election  as  aforesaid. 
"I  remain,  sir,  with  much  respect, 

"Your  most  obedient  servant, 

"JOHN  S.  WHITING, 
"Secretary  of  the  Medical  Society  of  the  Territory  of  Michigan." 

The  first  experiments  were  carried  on  at  Mackinac  and  were 
continued  at  Fort  Niagara,  to  which  place  Beaumont  was  removed. 
While  on  a  visit  to  Burlington,  Vt.,  as  one  of  his  master's  household, 
Alexis,  whose  interest  in  science  had  long  ago  reached  the  vanishing 
point,  ran  away  and  was  lost  to  his  benefactor  for  some  time.  This 
ungrateful  act  on  the  part  of  the  French-Canadian  proved  a  sore  dis- 
appointment to  our  '"'Backwoods  physiologist."  His  experiments  up 
to  this  time  were  to  estimate  the  length  of  time  required  for  the 
digestion  of  certain  kinds  of  food,  which  were  suspended  in  the  stom- 
ach by  means  of  silk  threads  and  withdrawn  from  time  to  time  to  note 
the  changes  in  the  substances.  He  found  that  food  would  digest  more 
quickly  in  the  stomach  than  when  mixed  with  gastric  juice  in  vitro. 

Four  years  after  St.  Martin's  unceremonious  departure,  Beau- 
mont got  in  communication  with  him.  In  the  meantime  Alexis  had 
married  and  became  the  father  of  two  children.  The  doctor  took 
hirn,  his  wife  and  two  children  into  his  own  home,  where  Alexis  did 
duty  as  a  common  servant  when  not  employed  for  purposes  of  experi- 
mentation. Beaumont's  laboratory  equipment  consisted  of  a  thermo- 
meter, a  few  open  mouthed  vials  and  a  sand  bag.     His  observations 


BEAUMONT  29 

were  made  with  a  true  spirit  of  inquiry  and  with  no  particular  hypo- 
thesis to  support.  Fifty-six  experiments  were  made  between  Dec. 
6th,  1829,  and  April  9th,  1831.  Alexis,  with  his  wife  and  family,  were 
permitted  to  return  home  to  Quebec  on  the  promise  to  appear  when 
again  wanted.  Beaumont  had  felt  that  he  had  accomplished  about  all 
he  was  able  in  his  researches  on  gastric  digestion,  and  he  longed  to 
go  to  Europe  a  year  and  take  St.  Martin  with  him,  that  the  work 
might  be  pursued  farther  by  more  competent  physiologic  chemists. 
The  brevity  of  his  furlough  precluded  the  idea  of  going  abroad  and 
instead  he  remained  in  Washington  with  Alexis  where  he  found  his 
surroundings  very  congenial.  Access  to  the  works  of  European 
physiologists  in  the  library  and  recognition  from  many  of  the  promi- 
nent men  at  the  capital  made  his  sojourn  pleasant. 

Between  Dec.  1st.,  1832  and  March  1st,  1833,  we  find  recorded 
116  experiments,  some  in  confirmation  of  what  had  been  done  before. 
He  tested  the  temperature  of  the  stomach  when  full,  when  fasting, 
when  exercising,  when  resting,  also  the  length  of  time  required  to 
digest  various  food  substances.  He  also  experimented  to  disprove 
the  old  theory  of  maceration  or  mechanical  trituration. 

Seeks  Assistance  of  Two  Leading  Scientists:  In  1833  Beaumont 
sought  the  assistance  of  two  of  the  leading  scientific  men  of  the 
United  States,  Robley  Dunglinson,  professor  of  physiology.  Univer- 
sity of  Virginia,  and  Benjamin  Silliman,  professor  of  chemistry  at 
Yale.  Thanks  to  Beaumont's  painstaking  and  methodical  nature,  the 
correspondence  between  the  two  and  himself  had  been  carefully  pre- 
served, and  it  constitutes  an  excellent  account  of  the  physiology  of 
the  period.  A  sample  of  gastric  juice  from  St.  Martin's  stomach  was 
sent  Dunglinson  for  analysis  with  the  request  to  convey  to  the  giver 
the  results  and  to  refrain  from  publishing  anything  that  would  antici- 
pate the  labors  of  Beaumont  himself.  He  is  assured  that  the  profes- 
sor has  but  one  desire  in  the  prosecution  of  his  profession,  by  teaching 
and  practice  to  benefit  his  fellow  men,  which  could  always  be  done 
with  due  credit  without  forestalling  his  coadjutors  in  the  field  of 
science,  or  arrogating  to  himself  merit  to  which  he  might  be  but  sec- 
ondarily entitled.  Dunglinson  found  the  sample  of  gastric  juice  to 
contain  "free  muriatic  and  acetic  acid  and  phosphates  and  murates 
with  bases  of  potassa,  soda,  magnesia  and  lime  and  animal  matter 
soluble  in  cold  but  not  in  hot  water." 

Professor  Silliman,  to  whom  a  bottle  of  gastric  juice  was  also 
submitted,  suggested  that  a  sample  be  sent  to  Professor  Berzelius,  of 
Stockholm,  Sweden,  "as  the  man  of  all  others  best  qualified  to  investi- 
gate a  subject  of  such  deep  interest  to  mankind."  Accordingly  a 
bottle  of  the  digestive  fluid  was  packed  for  shipment.  Beaumont's 
disappointment  may  be  imagined  when  it  was  known  that  the  parcel 
was  delayed  over  two  and  a  half  months.  This  he  learned  about  the 
time  he  was  patiently  awaiting  the  results  of  the  Swedish  professor's 
investigations.  In  the  meantime  Beaumont  had  received  a  letter  from 
Professor  Silliman  enclosing  an  abstract  of  a  portion  of  a  system  of 
chemistry  by  Berzelius,  important  as  presenting  a  clear  idea  of  the 
knowledge  of  the  physiology  of  digestion  at  that  time  (1833).  The 
communication  states,  among  other  things,  that  Prout,  Tiedeman  and 
Gmelin  gave  the  best  notions  on  the  subject  of  gastric  juice  and  ex- 


30  PATHFINDERS  OP  PHYSIOLOGY 

plained  the  contradictory  statements  of  other  authors ;  at  one  time  it 
was  said  to  be  very  fluid  clear  and  neutral  in  reaction ;  then  alkaline, 
then  acid.  Prout  in  1824  declared  the  gastric  juice  to  contain  free 
hydrochloric  or  muriatic  acid,  the  result  of  an  experiment  made  on 
the  contents  of  the  stomach  of  an  animal  killed  soon  after  eating. 
Gmelin  and  Tiedeman  also  established  the  presence  of  free  hydroch- 
loric acid.  The  fluid  of  the  empty  stomach  was  found  to  be  slightly 
acid,  sometimes  neutral  and  the  acidity  was  in  proportion  to  the  quan- 
tity, becoming  very  acid  when  food  had  been  swallowed.  According 
to  Gmelin  and  Tiedeman,  the  salts  of  gastric  juice  were  principally 
sodium  chloride  and  potassium  chloride  in  small  quantities,  hydro- 
chlorate  of  ammonia  and  a  little  sulphate  of  potassium.  The  com- 
munication concludes  with  the  assertion  that  "no  organ  for  the  special 
secretion  of  the  gastric  juice  has  yet  been  discovered." 

Berzelius'  Reply  Disappointing:  Through  Professor  Silliman, 
Beaumont  eventually  heard  from  Berzelius,  whose  letter  was  dated 
July,  1834.  The  communication  upon  which  such  great  expectations 
were  placed  was  wholly  disappointing.  It  was  in  the  main  an  apology 
for  the  writer's  inability  to  work  with  the  gastric  fluid  with  prospects 
of  results  of  any  value,  owing  to  the  time  which  had  elapsed  since 
its  secretion  and  its  arrival  at  his  laboratory,  to  the  possible  alteration 
on  account  of  summer  heat,  and  to  the  inadequate  quantity  received. 

Nothing  but  the  utmost  zeal  and  love  for  the  work  could  account 
for  the  persistence  with  which  Beaumont  pursued  his  researches.  He 
felt  not  only  the  handicap  of  inadequate  resources  and  facilities  for 
experimentation,  but  St.  Martin  was  a  source  of  canstant  annoyance 
to  him.  He  would  leave  his  master  and  benefactor,  often  absent  for 
several  years,  when  by  overtures  in  the  shape  of  money  he  would  be 
prevailed  upon  to  return  and  furnish  the  precious  fluid  for  his  mas- 
ter's investigation.  Beaumont's  lot  was  cast  at  a  time  when  it  was 
difficult,  almost  impossible,  to  obtain  government  grants  for  the  pro- 
motion of  education.  His  work,  therefore,  has  been  accomplished  al- 
most entirely  at  his  own  expense. 

Attains  Fame  Through  His  Stomach:  St.  Martin  lived  the  life 
of  the  French  Canadian  habitant  mostly  in  poverty,  though  physically 
he  was,  the  larger  part  of  his  life,  in  good  condition.  Nine  years  after 
his  notable  accident,  we  are  told,  he  took  his  family  in  an  open  canoe 
via  the  Mississippi,  passing  St.  Louis,  ascended  the  Ohio  River,  then 
crossed  the  state  of  Ohio  to  the  lakes  and  descended  the  Erie  and 
Ontario  and  the  River  St.  Lawrence  to  Montreal,  the  trip  consuming 
the  interval  from  March  to  June.  He  was  able  to  engage  in  manual 
labor  requiring  considerable  strength  and  endurance.  Perhaps  his  ex- 
treme poverty  is  due  to  lack  of  thrift  and  to  intemperance,  for  we  are 
told  that  he  indulged  immoderately  in  the  "glass  that  cheers." 

The  longevity  of  the  habitant  is  evidenced  in  St.  Martin,  for 
he  lived  twenty-eight  years  after  the  death  of  Beaumont.  St.  Mar- 
tin's death  occurred  in  his  eighty-third  year.  Sir  William  Osier,  at 
the  time,  (1880)  a  resident  of  Montreal,  reading  of  his  death,  wrote 
the  local  physician  and  parish  priest  urging  them  to  secure  for  him 
the  privilege  of  an  autopsy,  and  at  the  same  time  offering  a  goodly 
sum  for  the  stomach,  which  he  intended  to  place  in  the  Army  Medi- 
cal Museum  at  Washington,  but  his  entreaties  were  of  no  avail,  the 


BEAUMONT  31 

body  was  interred  eight  feet  below  the  surface  of  the  ground,  after 
being  detained  at  home  much  longer  than  the  usual  period,  so  that 
decomposition  setting  in,  might  baffle  the  doctors,  and  prevent  any 
attempts  at  resurrection. 

Beaumont  Resigned  From  Army:  William  Beaumont  resigned 
his  position  as  army  surgeon  in  1839.  He  continued,  however,  to 
attend  the  families  of  the  officers  at  St.  Louis,  where  he  made  his 
home.  Owing  to  the  distance  from  St.  Louis  of  his  successor,  who 
was  stationed  ten  miles  away,  he  presented  an  account  to  the  War 
Department  for  professional  services  covering  a  period  of  a  few 
months,  which  services  he  conceded  "irregular  and  informal,"  but 
"correct  and  just."  On  receipt  of  his  account  the  surgeon-general 
threatened  either  to  ignore  the  bill  or  to  deduct  the  amount  from 
the  salary  of  Beaumont's  successor.  The  manner  in  which  Beau- 
mont received  the  threat  showed  the  independent  nature  of  the  man. 
He  declared  the  surgeon-general's  view  at  "absurd  opinion,  con- 
tracted view,  narrow-minded  vindictive  spirit  and  petty  tyrannical 
disposition,"  of  the  "weak,  waspish  and  wilful  head  of  a  medical  de- 
partment," and  congratulated  himself  over  having  the  "privilege  of 
detesting  a  man,  the  motives  and  the  mind  from  which  such  egregius 
folly,  parsimony  and  injustice  could  emanate  and  be  promulgated." 
The  Surgeon-General  was,  however,  unyielding,  and  Beaumont's  claims 
were  unrecognized. 

Though  severed  from  the  War  Department,  he  still  had  a  very 
lucrative  practice,  and  what  is  above  any  monetary  consideration,  de- 
voted friends,  and  was  very  happy  in  his  domestic  relations.  The 
following  paragraph  quoted  in  Dr.  Myer's  Life  and  Letters  of  Beau- 
mont gives  a  splendid  estimate  of  his  character: 

"Dr.  Beaumont  possessed  great  firmness  and  determination  of  purpose. 
Difficulties  which  would  have  discouraged  most  men,  he  never  allowed  to  turn 
him  from  his  course.  These  he  did  not  attempt  to  evade  but  to  meet  and  overcome. 
He  possessed  more  than  any  man  I  ever  knew,  a  knowledge  almost  intuitive  of  hu- 
man character.  You  might  have  introduced  him  to  20  different  persons  in  a 
day,  all  strangers  to  him,  and  he  would  have  given  you  an  accurate  estimate  of 
the  character  of  each,  his  peculiar  traits,  disposition,  etc.  He  was  gifted  with 
strong  natural  powers  which,  working  upon  an  extensive  experience  in  life,  re- 
sulted in  a  species  of  natural  sagacity,  which  I  suppose  was  something  peculiar 
to  him  not  to  be  attained  by  any  course  of  study.  His  temperament  was  ardent 
but  never  got  the  better  of  his  instructed  and  disciplined  judgment,  and  when- 
ever or  however  employed,  he  always  adopted  the  most  judicious  means  of  ob- 
taining ends  that  were  always  honored.  In  the  sick  room  he  was  a  model 
of  patience  and  kindness;  his  intuitive  perceptions  guiding  a  pure  benevolence 
never  failed  to  inspire  confidence.  Thus,  he  belonged  to  that  class  of  physicians 
whose  very  presence  affords  nature  a  sensible  relief." 

He  died  on  April  25th,  1853.  His  death  was  considered  the  re- 
sult of  injuries  he  received  by  slipping  on  icy  steps  while  making  a 
professional  visit.  What  a  satisfaction  such  a  life  must  be,  and  the 
resignation  with  which  one  might  approach  the  infirmities  of  old  age 
and  one's  final  destiny.  And  indeed  a  few  months  before  the  end  he 
breathed  forth  this  beautifuly  symphony: 


32  PATHFINDERS  OF  PHYSIOLOGY 

"Myself  and  wife,  not  unlike  Jolin  Anderson  my  Jo,  liave  climbed  the  hill  o' 
life  togither,  and  mony  a  canty  day  we've  had  wi'  ane  anither.  But  now  we 
maun  totter  down  life's  ebbing  wane  in  peaceful  quiet  ease  and  compitence,  with 
just  so  much  selfishness  and  social  sympathy  as  to  be  satisfied  with  ourselves,  our 
children  and  friends,  caring  little  for  the  formalities,  follies  and  fashions  of  the 
present  age.  *  *  *  Come  when  it  may,  we  only  ask  God's  blessing  on  our 
frosted  brows  and  hand  in  hand  we  will  go  to  sleep  together," 

DR.  BEAUMONT'S  BOOK. 

I  am  fortunate  in  having  before  me  an  original  copy  of  Dr.  Beau- 
mont's work.  The  title  page  bears  the  following  description:  "Ex- 
periments and  Observations  on  the  Gastric  Juice  and  the  Physiology 
of  Digestion,  by  William  Beaumont,  M.  D.,  Surgeon  in  the  United 
States  Army.  Plattsburg.  Printed  by  F.  P.  Allen,  1833."  The  vol- 
ume is  dedicated  to  Joseph  Lovell,  M.  D.,  Surgeon  General  of  the 
United  States  Army.  The  work  comprises  280  pages,  122  of  which 
deal  with  "Preliminary  Remarks  on  the  Physiology  of  Digestion." 
The  remainder  deals  with  Experiments  and  Observations  on  the  Stom- 
ach of  Alexis  St.  Martin.  The  first  part  is  divided  into  seven  sections, 
as  follows:  1st,  Of  Ailment;  Section  two  of  Hunger  and  Thirst; 
Section  three  of  Satisfaction  and  Satiety;  Section  four  of 
Mastication,  Insalivation  and  Deglutition;  Section  five  of  Digestion 
by  Gastric  Juice;  Section  six  of  the  Appearance  of  the  Villous  Coat 
and  of  Motions  of  the  Stomach;  Section  seven  of  Chylification  and 
Uses  of  the  Bile  and  Pancreatic  Juice.  There  are  three  illustrations, 
consisting  of  crude  wood  cuts  of  the  gastric  fistulse.  The  typograph- 
ical appearance  of  the  work  should  be  onsidered  creditable  consider- 
ing the  printing  art  at  the  time.  The  conclusion  of  the  second  part  of 
the  work  contains  51  inferences  made  from  the  foregoing  experiments 
and  observations.     Of  these  I  shall  quote  a  few : 

That  digestion  is  facilitated  by  minuteness  of  division  and  tenderness  of  fibre 
and  retarded  by  the  opposite  qualities. 

That  the  quantity  of  food  generally  taken  is  more  than  the  wants  of  the 
system  require,  and  that  excess,  if  persevered  in,  generally  produces  not  only 
functional  aberration  but  disease  of  the  coats  of  the  stomach. 

That  bulk  as  well  as  nutriment  is  necessary  to  the  articles  of  diet. 

That  oily  food  is  diflicult  of  digestion,  though  it  contains  a  large  proportion 
of  the  nutrient  principles 

That  stimulating  condiments  are  injurious  to  the  healthy  stomach. 

That  the  use  of  ardent  spirits  always  produces  disease  of  the  stomach  if 
persevered  in. 

That  the  agent  of  chymification  is  the  gastric  juice,  which  acts  as  a  solvent 
of  food  and  alters  its  properties. 

That  the  action  of  gastric  juice  is  facilitated  by  the  warmth  and  motions  of 
the  stomach. 

That  it  coagulates  albumin  and  afterwards  dissolves  the  coagulum. 
ciples. 

That  the  gastric  juice  is  secreted  from  vessels  distinct  from  the  mucous 
follicles. 

That  bile  is  not  ordinarily  found  in  the  stomach  and  is  not  commonly  neces- 
sary for  the  digestion  of  food,  but  assists  in  the  digestion  of  oily  foods. 


BEAUMONT  33 

That  the  inner  coat  of  the  stomach  is  of  pale  pink  color,  varying  in  its  hues 
according  to  its  full  or  empty  state. 

That  the  motions  of  the  stomach  produce  a  constant  churning  of  its  contents 
and  admixture  of  the  food  and  gastric  juice. 

That  these  motions  are  in  two  directions,  transverse  and  longitudinal. 

Beaumont  failed,  however,  to  ascribe  any  digestive  function  to 
the  saUva.  He  maintained  that  food  finely  divided  placed  directly 
into  the  stomach  was  as  completely  digested  as  that  which  entered 
by  the  oesophageal  route. 

When  he  began  his  work  the  status  of  the  physiology  of 
digestion  had  been  very  well  described  by  William  Hunter;  "some 
physiologists  will  have  it  that  the  stomach  is  a  mill ;  others  that  it  is 
a  fermenting  vat ;  other  again  that  it  is  a  stew  pan ;  but  in  my  view 
of  the  matter  it  is  neither  a  mill,  a  fermenting  vat  nor  a  stew  pan,  but 
a  stomach,  gentlemen,  a  stomach."  When  William  Beaumont  com- 
pleted his  labors  there  was  a  marked  advance  in  knowledge  of  the 
digestive  process.  Among  the  most  important  results  of  his  worjj 
was  his  complete  and  accurate  description  of  the  gastric  juice,  which 
has  been  quoted  in  so  many  text  books  since  his  day. 

"Pure  gastric  juice  when  taken  directly  out  of  the  stomach  of  a  healthy  adult, 
unmixed  with  any  other  fluid,  save  a  portion  of  the  mucus  of  the  stomach  with 
which  it  is  most  commonly,  perhaps  always  combined,  is  a  clear,  transparent  fluid; 
inodorous;  a  little  saltish,  and  perceptibly  acid.  Its  taste,  when  applied  to  the 
tongue,  is  similar  to  mucilaginous  water,  slightly  acidulated  with  muriatic 
acid.  It  is  readily  diffusible  in  water,  wine  or  spirits;  slightly  effervescent  with 
alkalies,  and  is  an  effectual  solvent  of  the  materia  alimentaria;  it  possesses  the 
property  of  coagulating  albumin  in  an  imminent  degree;  it  is  a  powerful  anti- 
septic, checking  the  putrefaction  in  meat;  and  effectually  restorative  of  healthy 
action  when  applied  to  old  foetid  sores  and  foul  ulcerating  surfaces." 

His  work  confirmed  the  observation  of  Prout,  that  the  acid  con- 
tents of  the  gastric  secretion  was  hydrochloric.  He  recognized  the 
fact  that  the  elements  of  the  gastric  juice  and  the  mucus  of  the  stom- 
ach were  a  separate  secretion.  He  established  by  direct  observation 
the  marked  influence  of  mental  states  on  the  secretion  of  gastric  juice 
and  on  digestion.  His  was  the  first  comprehensive  and  thorough 
study  of  the  motions  of  the  stomach ;  and  to  quote  Osier :  "His  study 
of  the  digestibility  of  different  articles  of  diet  in  the  stomach  remains 
today  one  of  the  most  important  contributions  ever  made  to  practical 
dietetics." 

A  German  edition  of  the  work  was  issued  in  1834.  In  1838  Sir 
Andrew  Combe,  an  eminent  English  physician,  published  an  English 
edition  of  the  work,  so  as  to  give  it  greater  publicity  in  the  British 
Isles.  Probably  no  fairer  or  more  impartial  estimate  of  the  value 
of  Beaumont's  contribution  to  science  has  been  made  than  that  of 
Sir  Andrew  in  his  preface  to  the  British  edition.  Answering  the 
objection  that  Beaumont  had  made  no  original  discovery  in  the  phys- 
ology  of  digestion,  this  advocate  claims  that  by  "separating  the  truth 
clearly  and  unequivocally  from  the  numerous  errors  of  fact  and  opin- 
ion with  which  it  was  mixed  up,  and  thus  converting  into  certainties 
points  of  doctrine  in  regard  to  which  positive  proof  were  previously 
inaccessible,  he  has  given  to  what  was  doubtful  or  imperfectly  known 


34  PATHFINDERS   OF  PHYSIOLOGY 

a  fixed  and  positive  value  which  it  never  had  before,  and  which,  being 
once  obtained,  goes  far  to  furnish  us  with  a  clear  connected  and  con- 
sistent view  of  the  general  process  and  laws  of  digestion." 


CLAUDE   BERNARD,   PHYSIOLOGIST. 

1813—1878 


CHAPTER  IV. 


GLYCOGENIC     FUNCTION     OF     THE     LIVER— VASOMOTOR 
NERVES— CLAUDE  BERNARD. 

"For  a  man  to  be  an  investigator  of  the  first  order  two  gifts  are  perequisite 
it  is  not  merely  necessary  to  possess  a  well-ordered  and  what  we  may  term  a 
philosophic  imagination,  to  possess  a  mind  that  is  capable  of  balancing  phenomena, 
seeing  their  relationship  and  deducing  problems  that  have  to  be  solved  and  the  way 
in  which  to  solve  them;  there  must  be  something  more,  namely,  a  mechanical 
ability,  a  love  for  technique,  and  a  capacity  to  construct  and  manipulate  the  ap- 
propriate instruments.  This  is  particularly  necessary  in  connection  with  physio- 
logical resarch." — Adami 

The  real  life  of  every  notable  character  lies  in  the  story  of  his 
achievement,  rather  than  in  how  he  passed  his  days.  Human  interest, 
however,  loves  to  dwell  on  the  details  of  how  he  moved  among  his 
fellowmen  and  the  vicissitudes  that  befel  him  on  his  path  through 
life.  Often  in  the  lives  of  our  greatest  men  these  details  which  con- 
stitute the  human  touches  have  not  been  recorded.  Not  every  John- 
son has  his  Boswell,  and  we  must  content  ourselves  with  the  frag- 
mentary data  that  have  been  preserved.  Such  has  been  the  fate  of 
Claude  Bernard,  the  first  centenary  of  whose  birth  is  now  the  sub- 
ject of  commemoration. 

Early  Life  and  Education — Let  me  give  a  brief  summary  of  his 
life.  He  was  bom  on  July  12th,  1813,  of  humble  parentage ;  his  father 
owned  a  small  farm  at  St.  Julien,  near  Lyons,  France.  The  vintage  of 
the  little  estate  which  was  situated  in  the  wine  district  of  France,  pro- 
vided the  family  revenue.  The  property  eventually  came  into  the 
hands  of  the  son,  who  spent  his  summers  there  within  view,  on  clear 
days,  of  the  white  summits  of  the  Alps.  Bernard  received  his  early 
education  at  his  native  village  and  afterwards  at  Lyons.  His  educa- 
tion was,  however,  cut  short  by  necessity,  which  turned  him  to  prac- 
tical pharmacy  as  a  means  of  earning  a  living.  The  young  man  pos- 
sessed that  "fine  frenzy"  which  makes  "the  lunatic,  the  lover  and  the 
poet"  of  "imagination  all  compact,"  and  was  on  the  point  of  giving  up 
the  calling  which  had  engaged  his  attention  for  two  years,  for  litera- 
ture. His  literary  aspirations  drew  him  towards  the  dramatic  art,  and 
it  is  hard  to  predict  what  the  future  physiologist  might  have  given  to 
the  world  had  not  the  divine  flame  been  smothered  by  a  more  prosaic 
career  of  investigator.  He  was  the  author  of  a  comedy,  "The  Rose  of 
the  Rhone,"  which  had  met  with  a  certain  amount  of  success.  But  des- 
tiny had  reserved  Bernard  for  another  and  very  different  calling. 
He  submitted  his  work  to  the  great  French  critic,  St.  Marc  Girardin, 
who,  while  recognizing  its  merit,  advised  the  young  aspirant  to  lit- 
erary fame  to  pursue  a  more  lucrative  calling,  to  engage  in  some 
pursuit  in  which  he  could  earn  his  bread  and  to  court  the  Muses  only 
in  his  leisure  moments.  "You  have  studied  pharmacy,"  said  the 
critic,  "study  medicine;  you  will  thereby  much  more  surely  gain  a 
livelihood."    Bernard  followed  this  advice  with  heart  and  soul,  de- 


36  PATHFINDERS  OF  PHYSIOLOGY 

fraying  his  expenses  by  tutorage.  The  Hterary  longings  began  to 
fade  as  the  young  savant  waxed  warm  with  his  medical  studies. 
Anatomy  and  physiology  claimed  the  greater  portion  of  his  attention 
and  energy.  His  remarkable  manual  dexterity,  in  which  he  was  par- 
ticularly fortunate,  rendered  his  dissections  of  singular  completeness 
and  value.  The  chaotic  condition  of  physiology  of  the  time  (1840) 
served  to  awaken  in  his  mind  a  desire  to  solve  problems  by  direct 
experimental  appeal  to  nature.  He  was  one  of  the  first  to  employ 
animal  experimentation,  or  vivisection.  In  1841  he  attracted  the 
attention  of  the  great  Majendie,  then  the  leading  physiologist  of 
France,  also  Professor  of  Medicine  in  the  College  of  France.  Ma- 
jendie is  described  as  being  in  manner  abrupt  and  even  rough  and 
rude.  At  first  he  took  little  notice  of  Bernard,  his  new  interne,  but 
was  soon  impressed  with  the  young  man's  dexterity  and  skill.  One 
day  while  Bernard  was  busy  at  his  dissecting,  Majendie  blurted  out: 
"I  say,  you,  there.  I  take  you  as  my  preparateur  at  the  College  of 
France."  And  it  was  not  long  before  the  master  had  occasion  to  say 
in  his  gruff  way  as  he  left  the  class-room:  "You  are  a  better  man 
than  I  am."  Bernard's  career  as  physiologist  may  be  said  to  date 
from  this  appointment  in  1841. 

Claude  Bernard  was  of  a  retiring,  silent  nature,  difficult  to  under- 
stand and  often  misunderstood.  Michael  Foster  described  him  as 
"tall  in  stature,  with  a  fine  presence  and  a  noble  head,  the  eyes  full 
at  once  of  thought  and  kindness ;  he  drew  the  look  of  observers  upon 
him  wherever  he  appeared.  As  he  walked  the  streets  passers-by 
might  be  heard  to  say,  *I  wonder  who  that  is ;  he  must  be  some  dis- 
tinguished man.' " 

The  Productive  Period — Bernard  had  shown  the  precious  metal  of 
his  genius  before  he  was  far  on  in  his  twenties.  Nearly  all  of  his  great 
achievements  were  accomplished  during  the  period  of  his  life  which 
ended  with  1860.  The  essential  results  of  his  two  greatest  discoveries, 
the  glycogenic  function  of  the  liver  and  the  vaso  motor  nerves  were 
gained  prior  to  1850,  before  he  was  37  years  old.  He  is  illustrative  of 
Osier's  declaration  that  the  world's  best  and  most  important  work 
was  mainly  done  by  young  men,  for  further  example:  Mor- 
gagni's  germinal  idea,  which  made  him  the  father  of  modem  path- 
ology, came  to  him  when  he  was  scarcely  twenty;  Auenbrugger 
began  his  work  upon  percussion  when  he  was  under  twenty-five ;  Laen- 
nec  undertook  the  problem  of  constructing  a  system  of  auscultation  in 
his  early  twenties  and  published  his  book  when  he  was  not  yet  thirty- 
five. 

All  significant  work  in  medicine  has  had  its  basis  in  observation, 
not  theory.  Men  have  been  prone  to  theorize  too  much  and  to  observe 
too  little.  For  two  thousand  years  the  learned  men  of  Europe  de- 
bated as  to  whether  this  or  that  place  was  the  site  of  ancient  Troy,  or 
whether  there  ever  was  such  a  place  at  all.  It  remained,  however,  for 
a  retired  man  of  business,  Schliemann,  to  decide  the  question.  He 
said,  "Let  us  go  and  see,"  and,  at  the  expense  of  a  few  thousand 
pounds,  he  went  and  found  Troy  and  Mycenae  and  revealed  or  dis- 
covered the  whole  matter — "The  most  tremendous  and  picturesque  tri- 
umph of  the  scientific  method  over  mere  talk  and  pretended  historic 
learning,"  says  Ray  Lancaster,  "which  has  ever  been  since  human 


GLYCOGENIC  FUNCTION  OP  LIVER  37 

record  has  existed."  Emerson  has  said:  "I  am  impressed  with  the 
fact  that  the  greatest  thing  a  human  soul  ever  does  in  this  world  is 
to  see  something  and  tell  what  it  saw  in  a  plain  way.  Hundreds  of 
people  can  talk  for  one  who  can  think,  but  thousands  can  think  for 
one  who  can  see.  To  see  clearly  is  poetry,  philosophy  and  religion  all 
in  one."  And  we  might  add,  that  rare  quality  of  mind  which  enables 
its  possessor  to  see  clearly  is  the  sine  quo  non  of  the  true  scientist. 

Gastric  Digestion — Among  Bernard's  earliest  investigations  was 
that  of  gastric  digestion.  It  was  important  chiefly  as  a  prelude  to  the 
momentous  discoveries  he  afterwards  made.  He  was  the  first  to  in- 
quire into  the  differences  to  be  found  between  the  digestive  appara- 
tuses and  functions  of  plant-eating  and  meat-eating  animals — between 
the  herbivora  and  carnivora.  The  former  thoroughly  masticate  their 
food,  while  the  latter  bolt  theirs.  This  instinct  is  explained  by  the  fact 
that  the  food  of  the  plant-eating  animal  contains  a  relatively  large 
amount  of  starch,  requiring  thorough  admixture  of  saliva  as  an  aid  to 
its  digestion.  Those  animals  subsisting  on  meat-protein  do  not  require 
the  aid  of  the  saliva,  which  accounts  for  the  rapidity  with  which  they 
devour  their  food.  From  this  Bernard  turned  to  study  the  function 
of  the  pancreatic  juice.  Up  to  this  time  the  pancreas  had  been  passed 
over  in  silence  by  the  physiologists  of  the  day.  He  demonstrated  its 
three-fold  action:  "He  showed  that  it,  on  the  one  hand,  emulsified, 
and,  on  the  other  hand  split  up  into  fatty  acids  and  glycerine,  the 
neutral  fats  that  are  discharged  from  the  stomach  into  the  duodenum. 
He  proved  it  had  a  powerful  action  on  starch,  converting  it  into 
sugar."  The  study  of  the  action  of  the  pancreatic  juice  upon  proteins 
begun  by  Bernard  was  continued  by  Kuhne,  his  pupil,  who  investi- 
gated the  action  of  extracts  of  the  gland.  Pancreatic  juice  as  se- 
creted does  not  possess  proteolytic  powers.  This  change  under  normal 
conditions  is  brought  about  by  the  activating  substance,  enteroki- 
nase,  contained  in  the  succus  entericus  producing  as  soon  as  the 
pancreatic  juice  enters  the  gut,  the  change  from  the  inert  typsinogen 
to  trypsin,  thus  acquiring  an  activity  over  proteins  superior  to  that 
of  any  other  digestive  juice.  (Starling).  Up  to  Bernard's  time  the 
principal  role  of  digestion  had  been  confined  to  the  gastric  juice.  With 
his  discoveries  it  became  clear  that  the  action  of  the  gastric  juice  on 
the  food  in  the  stomach  was  simply  preliminary  to  intestinal  diges- 
tion and  that  the  chief  work  in  the  preparation  of  the  food  for  ab- 
sorption was  accomplished  by  the  pancreatic  juice. 

Discovers  Glycogenic  Function  of  Liver — Important  as  were  his 
numerous  contributions  to  our  knowledge  of  physiology,  Claude  Ber- 
nard is  probably  best  known  as  discoverer  of  the  glycogenic  function 
of  the  liver.  The  story  of  his  discovery  is  interesting  and  well  worth 
relating.  The  dominant  opinion  among  physiologists  when  Bernard  be- 
gan his  work  was  to  the  effect  that  animals  and  plants  presented  a 
chemical  contrast  to  each  other.  The  plant  built  up  such  organic  com- 
pounds as  fats,  carbohydrates  and  proteins  out  of  inorganic  elements ; 
the  animal  feeding  on  the  plant  received  these  organic  compounds  into 
its  body  resolving  them  into  inorganic  substances,  at  the  same  time 
using  that  resolution  for  the  needs  of  life.  While  the  animal  modified 
vegetable  proteins,  carbohydrates  and  fats  so  as  to  give  them  an  ani- 
mal character,  it  never  made  anything  new.    It  was  maintained  that 


38  PATHFINDER  OF  PHYSIOLOGY 

the  animal  body  never  manufactured  any  of  these  three  compounds, 
that  all  or  any  of  them  present  in  the  animal  body  had  been  taken  into 
it  with  its  food. 

Such  was  the  current  belief  among  physiologists  of  France  at 
the  beginning  of  the  fourth  decade  of  last  century.  The  first  heresy 
was  uttered  by  Liebig  who  proved  that  the  fat  accumulated  in  the 
bodies  of  fattened  geese  exceeded  greatly  the  quantity  of  fat  in  the 
intake  of  food,  and  furthermore  that  when  a  cow  was  fattened,  the 
excreta  during  the  fattening  period  contained  as  much  fat  as  the  food 
taken.  At  this  time  Bernard  undertook  his  researches  on  the  physi- 
ology of  sugar.  His  first  discovery  was  that  cane  sugar  acted  upon 
by  the  gastric  juice  was  changed  into  dextrose  (glucose).  It  was  his 
intention  to  study  the  three  great  classes  of  foods,  but  he  found  it  nec- 
essary to  confine  his  attention  to  the  carbohydrates  owing  to  the  fas- 
cinating problems  suggested  by  diabetes.  He  set  about  to  discover  the 
cause  of  the  excess  of  sugar  in  diabetes  with  the  hope  of  finding  a 
remedy  for  the  disease. 

Having  previously  satisfied  himself  that  no  dextrose  was  present 
in  the  alimentary  canal,  or  in  the  portal  blood,  Bernard  fed  a  dog  on 
meat  only;  killing  the  animal  at  the  height  of  digestion  he  found  to 
his  great  astonishment  the  blood  loaded  with  dextrose. 

"Why!"  said  he,  "if  I  have  made  no  mistakes  I  have  in  this  ex- 
periment come  upon  the  production  of  sugar ;  the  liver  produces  sugar. 
If  the  result  I  have  got  is  confirmed  on  repetition  of  the  experiment, 
the  liver  is  the  sugar-producing  tissue.  It  manufactures  sugar  out 
of  something  that  is  not  sugar,  and  within  it  lies  the  secret  of  dia- 
betes. This  is  a  big  thing  of  which  I  have  got  hold,  I  must  make 
sure  that  I  have  made  no  mistake  in  the  experiment,  and  then  push 
forward  as  far  as  possible  the  lead  thus  given  me." 

Bernard's  results  were  confirmed  by  numerous  experiments.  He 
determined  that  the  sugar  in  question  was  dextrose,  responding  to  all 
the  tests  for  dextrose.  He  also  discovered  that  while  this  hepatic 
sugar  did  not  come  direct  from  the  food,  it  was  influenced  in  regard 
to  its  quantity  by  the  nature  of  the  food.  Starling,  however,  main- 
tains that  in  some  animals,  the  carnivora,  the  liver  can  continue  to 
supply  sugar  to  the  blood  on  a  diet  which  includes  only  proteins  and 
fats.  Von  Noorden  explains  the  fact  that  proteins  yield  sugar,  by 
the  presence  of  a  carbohydrate  group  in  the  protein  molecule,  which  is 
split  off  during  pepsin-hydrochloric  acid  digestion. 

Bernard  eventually  came  to  the  conclusion  that  sugar  was  not 
formed  immediately  from  the  elements  whatever  they  raight  be  which 
the  blood  brought  to  the  liver,  but  from  some  substance  existing  in 
the  fiver  tissue  which  was  capable  of  being  converted  into  sugar.  In 
1857  he  announced  to  the  scientific  world  the  discovery  of  glycogen. 
Though  he  made  known  each  step  in  his  discoveries  which  extended 
over  a  number  of  years,  he  had  the  satisfaction  of  telling  the  whole 
story  in  his  own  writings,  never  having  experienced  the  humiliation 
which  is  sometimes  the  lot  of  pioneers,  in  seeing  their  leading  con- 
ceptions worked  out  by  other  minds.  To  quote  his  biographer,  Sir 
Michael  Foster,  "Bernard  in  the  matter  of  glycogen  not  only  laid  the 
first  stone  but  left  a  house  so  nearly  finished  that  other  men  have  been 
able  to  add  but  little." 


VASO-MOTOR  NERVES  39 

"No  less  pregnant  of  future  discoveries,"  says  this  biographer, 
"was  the  idea  suggested  by  this  newly  found-out  action  of  the  hepatic 
tissue,  the  idea  happily  formulated  by  Bernard  as  'internal  secretion.' 
No  part  of  physiology  is  at  the  present  day  being  more  fruitfully 
studied  than  that  which  deals  with  the  changes  the  blood  undergoes 
as  it  sweeps  through  the  several  tissues."  The  study  of  these  in- 
ternal secretions  constitutes  a  path  of  inquiry  which  has  within  re- 
cent years  been  pursued  with  conspicuous  success. 

To  Bernard  we  owe  the  discovery  of  the  remarkable  fact  that 
temporary  diabetes  may  be  caused  by  puncture  of  the  fourth  ven- 
tricle. This  glycosuria  was  formerly  attributed  to  direct  stimula- 
tion of  the  liver  through  its  nervous  connections.  It  has  been  found, 
however,  that  if  the  left  adrenal  is  cut  off  from  the  left  sympathetic 
nerve,  no  sugar  appears  in  the  urine  after  the  medulla  has  been  punc- 
tured, and  it  is  now  believed  that  the  stimulus  is  transmitted  by  the 
left  sympathetic  nerve  to  the  left  adrenal,  whence  it  is  passed  to  the 
right  adrenal  by  the  connecting  nerves.  As  a  consequence  of  the 
medullary  puncture  the  adrenals  secrete  more  actively  and  the  in- 
creased flow  of  the  adrenal  secretion  in  its  turn  brings  about  an 
excessive  output  of  sugar  by  the  liver.*  A  number  of  toxic  influences 
possibly  act  in  the  same  way,  the  glycosuria  to  which  they  give  rise 
being  partly  the  result  of  the  action  they  exert  on  the  diabetic  center 
in  the  medulla,  and  partly  an  effect  of  their  stimulating  action  on  the 
sympathetic  nerves,  or  on  the  adrenals  directly,  thus,  in  any  case, 
causing  hyperfunction  of  the  chromaffin  system,  with  consequent 
overproduction  of  sugar  by  the  liver. 

Discovery  of  Vaso-Motor  Nerves — Next  in  importance  to  the  dis- 
covery of  glycogen  was  Bernard's  discovery  of  the  vaso-motor 
nerves.  "To  Claude  Bernard,"  says  Sir  Michael  Foster,  "we  owe  the 
foundations  of  the  vaso-motor  system.  He  made  known  to  us  the  ex- 
istence of  vaso-motor  nerves  and  he  also  made  known  to  us  that  vaso- 
motor nerves  are  of  two  kinds,  vaso-constrictor  and  vaso-dilator — ^the 
two  fundamental  facts  of  vaso-motor  physiology."  The  importance 
of  this  discovery  can  hardly  be  over-estimated  when  we  consider  that 
there  is  scarcely  a  physiological  problem  of  any  magnitude  which  does 
not  sooner  or  later  involve  vaso-motor  questions.  The  vaso-motor 
nerves  presiding  as  they  do  over  the  contraction  and  dilation  of  the 
walls  of  the  blood  vessels,  assume  an  important  role  in  such  functions 
as  gastric  digestion,  blood  pressure,  heat  processes,  blushing  and 
various  other  congestions,  or  on  the  other  hand,  the  significant  blanch- 
ing of  an  organ  as  in  sudden  fright. 

Among  Bernard's  minor  investigations  which  might  be  mentioned 
is  that,  into  the  physiological  action  of  curare,  a  black  resenoid  ex- 
tract prepared  by  the  South  American  Indians  from  the  bark  of  strych- 
nos  toxifera  and  used  to  poison  arrows.  Owing  to  its  poor  diffusi- 
bility  through  animal  membranes  curare  i^  harmless  taken  into  the 
alimentary  canal,  though  the  minutest  quantity  introduced  into  a 
wound  is  fatal.,  Since  Bernard's  time  curare  has  become  an  instru- 
ment in  the  hands  of  the  physiologist  to  enable  him  to  abolish  tempo- 
rarily the  movements  of  the  skeletal  muscles,  enabling  him  to  carry 
out  experiments  which  could  not  be  made  without  such  aid. 

The  precise  action  of  carbonmonoxide  gas  in  asphyxia  no  one 
understood  until  Bernard  investigated  the  matter.    His  experiments 

♦Futcher  Journal  A,  M.  A.  December  21,  1912. 


40  PATHFINDERS  OF  PHYSIOLOGY 

led  him  to  conclude  that  C  0  was  rapidly  poisonous  to  animals  owing 
to  the  fact  that  it  instantly  displaced  the  oxygen  of  the  red  corpuscle 
and  could  not  itself  be  subsequently  displaced  by  oxygen.  The  animal 
died  because  the  red  corpuscles  were,  so  to  speak,  paralyzed  and  cir- 
culated as  inert  bodies  devoid  of  the  power  of  sustaining  life. 

A  Friend  of  Pasteur — It  is  interesting  to  note  that  at  a  time 
when  physiological  opinion  favored  spontaneous  generation,  vitalism 
and  such  theories,  the  independent  mind  of  Claude  Bernard  foresaw 
what  subsequent  decades  of  physiological  research  have  found  to  ap- 
proximate the  truth  on  such  subjects.  He  was  a  firm  friend  of  Pas- 
teur, whom  he  ably  seconded  in  his  efforts  to  disprove  spontaneous 
generation. 

A  man  is  great  in  proportion  to  the  obstacles  he  is  able  to  sur- 
mount. The  subject  of  this  paper  illustrates  the  truth  that  one  who 
possesses  in  a  high  degree  the  qualities  of  genius  will  succeed  in  spite 
of  his  surroundings.  His  early  education,  neither  adequate  nor  con- 
ducive of  the  best,  together  with  the  keen  struggle  for  a  livelihood, 
and  in  his  early  career,  the  apathy  of  an  unappreciative  age  and  labor- 
atories with  meagre  equipment,  were  obstacles  which  bring  into  relief 
the  rare  qualities  that  he  possessed.  Contrast  such  a  condition  with 
the  magnificent  equipment  and  endowment  of  modern  scientific  re- 
search and  the  facilities  for  training  as  they  exist  today ! 

Bernard's  life  was  far  from  being  strewn  with  roses.  He  was  mar- 
ried to  a  wife  who  was  non-appreciative  of  his  genius.  She  saw  noth- 
ing in  what  to  her  was  empty  honor,  the  homage  of  the  scientific 
world,  when  the  means  which  make  for  affluence  were  not  forthcom- 
ing. His  two  daughters  became  estranged  from  him  and  it  is  said 
that  one  of  them  who  was  still  living  within  the  last  ten  years,  joined 
that  silly  sentimental  class  of  antivivisectionists  and  endowed  hos- 
pitals for  dogs  and  cats  to  atone  for  the  crimes  of  vivisection  which 
her  father  had  committed.  Not  only  lacked  he  the  sympathy  which 
"in  true  marriage  lies,"  but  he  began  his  work  at  a  time  when  the 
physiologist  had  need  of  a  "real  passion  for  his  science  and  in  order  to 
ward  off  fatal  discouragement  had  to  possess  his  soul  of  high  courage 
and  great  patience.  So  soon  as  the  experimental  physiologist  was  dis- 
covered he  was  denounced ;  he  was  given  over  to  the  reproaches  of  his 
neighborhood  and  subjected  to  the  annoyance  of  the  police;"  Bernard 
suffered  all  this. 

But  conscientious  work  well  performed  is  not  without  its  rewards 
and  perhaps  the  greatest  is  the  satisfaction  of  "something  attempted, 
something  done."  He  was  a  greater  man  than  Majendie,  whose  re- 
searches were  made  more  or  less  at  random  and  who  had  described 
himself  as  a  "rag  picker  by  the  dust  heap  of  science."  Bernard  always 
made  his  experiments  with  a  definiteness  of  purpose.  His  contribu- 
tions to  physiology  have  been  greater  in  number  and  importance  than 
those  of  any  other  investigator.  Later  in  life  he  enlisted  the  friend- 
ship of  Emperor  Napoleon  HI.,  which  resulted  in  two  well  equipped 
laboratories  which  greatly  facilitated  his  work.  His  academic  oppor- 
tunities included  professorships  in  the  College  of  France  as  well  as  a 
chair  at  the  Sorbonne.  In  1868,  he  was  admitted  to  the  Academy  of 
France  and  made  one  of  the  "Immortals." 


SKILL  AS  EXPERIMENTER  41 

The  Quest  for  Truth — As  already  mentioned,  Bernard  possessed 
a  faculty  that  contributed  in  no  small  degree  to  his  success  as  phy- 
siologist. Huxley  has  described  an  educated  man  as  one  whose  hand 
is  the  ready  servant  of  his  will.  Often,  too,  great  stress  is  laid  upon 
the  purely  intellectual  qualities  and  too  little  upon  that  manual  dex- 
terity which  is  so  essential  to  successful  work  in  the  laboratory. 
In  fact  medicine  itself  is  an  art  as  well  as  an  ensemble  of  sciences, 
and  the  art  is  as  important  as  the  science.  As  much  depends  upon 
the  skillful  use  of  the  senses,  and  in  surgery,  skill  in  manipulation, 
as  upon  the  well  trained  mind.  The  extreme  nicety  with  which  Ber- 
nard performed  his  dissections  excited  the  astonishment  as  well  as 
the  admiration  of  his  associates.  It  was  this  faculty  which  first  won 
him  the  favor  of  Majendie.  A  clumsy  experiment  is  apt  to  be  a  poor 
experiment  barren  of  results,  and  a  patient's  chances  of  life  may  be 
jeopardized  by  an  operation  poorly  performed. 

Bernard  was  active  until  the  end.  On  what  proved  to  be  his  death- 
bed he  worked  at  the  revision  of  proofs  of  a  volume  of  lectures  on 
operative  physiology.  He  died  on  the  tenth  of  February,  1878,  and 
was  laid  in  the  grave  with  all  the  pomp  and  ceremony  of  a  state  fu- 
neral. Gambetta  eulogized  him  as  one  who  had  never  allowed  himself 
to  be  led  away  either  by  party  spirit  or  by  the  dogmas  of  a  school,  or 
by  private  feelings.  Bernard's  work  is  a  model  of  patient  persevering 
investigation,  experiment  and  research,  an  unprejudiced  and  disinter- 
ested quest  for  truth.  He  lived  up  to  and  fulfilled  the  ideals  with 
which  he  began  his  career,  ideals  aptly  expressed :  "Truth  like  beauty 
is  when  unadorned,  adorned  the  most."  Such  ideals  have  inspired  men 
of  light  and  leading  of  all  time;  they  inspire  medicine  today,  ideals 
old  yet  always  new,  and  we  may  say  with  Kipling : 

"The  men  bulk  big  on  the  old  trail,  our  own  trail,  the  out  trail. 
They're  God's  own  guides  on  the  Long  Trail,  the  trail  that's  always  new. ' 


CHAPTER  V. 


RESPIRATION 

The  ancients  speculated  upon  the  physiology  of  respiration; 
Aristotle  (384  B.  C.)  contended  that  the  function  of  breathing  was  to 
cool  the  blood.  It  was  noticed  that  animals  over-heated  from  exertion 
breathed  more  rapidly,  hence  the  inference.  Galen  (131-203  A.  D.) 
also  maintained  that  the  air  inspired  served  to  regulate  and  to  cool 
down  the  innate  heat  of  the  heart;  that  the  peculiar  action  of  the 
chest  wall  seen  in  respiration  introduced  into  the  blood  the  air  re- 
quired for  the  regeneration  of  vital  spirits  in  the  left  side  of  the 
heart,  whence  by  the  arterial  route  they  were  distributed  through- 
out the  body.  Galen  also  recognized  the  necessity  of  ridding  the  body 
of  "fulginous  vapors*'  produced  by  the  innate  fire  in  the  heart  which 
act  was  accomplished  by  expiration.  In  the  latter  part  of  the  fifteenth 
century,  Leonardo  da  Vinci,  painter,  mathematician  and  naturalist, 
disproved  the  fallacy  that  air  simply  cooled  the  blood  in  respiration. 
He  found  that  air  was  consumed  by  fire  and  that  animals  could  not 
live  in  a  medium  incapable  of  supporting  combustion.  This  is  the 
first  record  in  the  history  of  science  which  pointed  to  the  fact  that 
the  function  of  air  in  respiration  depended  upon  its  chemical  com- 
position and  not  upon  its  physical  properties. 

It  is  evident  that  no  real  advance  could  be  made  in  the  physiology 
of  breathing  until  the  circulation  of  the  blood  had  been  demonstrated. 
Furthermore,  this  department  of  the  science  of  physiology  lagged 
until  the  chemist  appeared  on  the  scene.  Harvey  had  pointed  out 
that  as  the  blood  went  to  the  lungs  from  the  right  side  of  the  heart 
thence  to  the  left  auricle  a  marked  change  took  place,  the  blood  as- 
suming a  bright  arterial  hue.  The  cause  which  resulted  in  this 
peculiar  change,  Harvey  was  unable  to  discern,  nor  did  it  become 
known  until  a  much  later  day,  when  scientists  became  familiar  with 
the  characteristics  and  constituents  of  atmospheric  air. 

Mechanics  of  Respiration.  The  first  real  knowledge  on 
the  mechanics  of  respiration  we  owe  to  Borelli.  Applying 
the  knowledge  of  muscular  contraction  on  the  one  hand, 
and  atmospheric  pressure  on  the  other,  he  taught  that  inspira- 
tion consisted  of  the  entrance  of  air  into  the  chest  by  virtue  of  at- 
mospheric pressure,  the  thorax  being  enlarged  by  the  muscular  con- 
traction of  its  walls;  expiration  consisted  mainly  in  a  cessation  of 
muscular  contraction.  Borelli  broke  with  the  ancient  view  that  the 
function  of  breathing  was  the  cooling  of  the  excessive  heat  of  the 
heart  or  the  ventilation  of  the  vital  flame.  "So  great  a  machinery 
and  vessels  and  organs  of  the  lungs,"  he  continues,  "must  have  been 
instituted  for  some  grand  purpose ;  and  that  we  will  try  to  expound, 
if  possible,  though  we  shall  stammer  as  we  go  along."  Again  he  in- 
sists, "Air  taken  in  by  breathing  is  the  chief  cause  of  the  life  of  ani- 
mals." It  is  more  important  than  the  heart  and  the  circulation  of  the 
blood. 


ROBERT  BOYLE  43 

The  Work  of  Boyle.  Now  we  turn  to  the  Eng-lish 
school.  Robert  Boyle  (1627-1691),  perhaps  the  most  re- 
nowned physicist  of  his  time,  by  means  of  the  air  pump 
made  many  researches  on  the  "spring"  of  air.  He  showed, 
among  other  things  that  a  flame  was  extinguished  in  a  partial 
vacuum  and  that  in  a  more  complete  vacuum  not  only  the  flame  but 
the  lives  of  small  animals  such  as  the  mouse  ceased  very  quickly. 
Here  we  see  that  the  phenomena  connected  with  the  burning  candle 
closely  resembled  the  phenomena  of  life ;  furthermore  that  air  what- 
ever it  might  be,  and  not  the  mechanical  movements  of  the  chest 
wall  was  necessary  for  the  continuance  of  life.  Boyle  lived  at  Ox- 
ford for  many  years  and  while  there  made  important  improvements 
in  the  air  pump  and  in  a  long  series  of  experiments  with  it  made  vari- 
ous discoveries  in  the  properties  of  air  and  the  propagation  of  sound. 
He  was  at  the  same  time  an  ardent  student  of  theology.  He  was  ad- 
vised to  enter  the  church,  but  declined,  feeling  that  his  writings  on 
religious  topics  would  have  greater  weight  coming  from  a  layman 
than  from  a  paid  clergyman.  As  a  man  of  science  he  was  the  first 
to  carry  out  the  principles  of  Bacon's  Novum  Organum. 

The  next  step  was  taken  by  Robert  Hooke,  who  was  for  some 
time  assistant  to  Boyle.  Hooke  was  born  on  the  Isle  of  Wight,  in  1635. 
Hg,p»vas  destined  for  the  church,  but  ill-health  diverted  his  career  into 
other  channels,  which  gave  scope  for  his  precocious  mechanical 
•genius.  His  personal  appearance  is  described  as  very  unattractive; 
his  hair  being  in  dishevelled  locks  over  his  haggared  countenance.  He 
possessed  an  irritable  temper  and  was  much  given  to  spending  his 
time  in  solitude. 

To  him  Boyle  was  endebted  for  valuable  work  in  connection  with 
the  perfecting  of  his  air  pump.  He  was  one  of  the  earliest  and  most 
zealous  users  of  the  microscope ;  a  volume  entitled  Micrographia,  con- 
tains an  account  of  his  many  "Observations  Made  on  Minute  Bodies 
of  varied  kinds  by  magnifying  glasses."  Hooke's  microscopic  studies 
on  cork  lead  to  the  adoption  of  the  term  "cell"  as  the  histologic  unit. 
He  was  curator  of  the  Royal  society,  at  a  meeting  of  which  he  demon- 
strated before  the  Fellows  an  experiment  on  artificial  respiration, 
which  had  been  made  before  and  many  times  since.  The  uniqueness 
of  the  experiment  consisted  in  the  important  conclusions  which  Hooke 
made.  The  experiment  consisted  in  opening  the  thorax  of  a  dog  and 
substituting  the  movements  of  the  chest  wall  by  respiratory  move- 
ments accomplished  by  means  of  hand  bellows,  the  nozzel  inserted  in 
the  trachea.  This  proved  that  the  mechanical  movements  of  the 
chest  wall  were  only  of  a  secondary  importance  and  that  the  whole 
business  of  respiration  was  carried  on  in  the  lungs.  This  fact  was  fur- 
ther proven  by  inflating  the  lungs  to  their  utmost  capacity  and  keep- 
ing them  distended  by  a  powerful  blast  allowing  the  air  to  escape  con- 
tinually through  minute  holes  pricked  in  the  lungs.  This  showed  that 
life  could  be  maintained  even  in  the  absence  of  the  artificial  move- 
ments so  long  as  the  parenchyma  of  the  lungs  were  so  subjected  to  a 
fresh  supply  of  air.  Therefore  the  secret  of  the  change  from  venous 
to  arterial  blood  depended  upon  the  exposure  of  the  blood  to  fresh 
air  which  was  in  the  course  of  life  accompHshed  by  the  bellows-like 
action  of  the  chest  wall  and  diaphragm. 


44  PATHFINDERS  OF  PHYSIOLOGY 

Change  in  Color  of  Venous  to  Arterial  Blood. — Richard  Lower, 
1631,  concluded  that  the  change  in  color,  venous  to  arterial,  blood  was 
due  to  the  exposure  of  the  blood  to  the  air  in  the  lungs ;  he  drew  the 
further  conclusion  that  the  change  in  color  was  due  not  to  the  ex- 
posure alone,  but  to  the  fact  that  the  blood  took  up  some  of  the  air; 
that  is,  according  to  Lower,  arterial  blood  differed  from  venous  in 
that  it  contains  air.  The  blood  gave  up  its  "fresh  air"  in  the  course 
of  the  circulation,  hence  the  necessity  of  a  constant  supply  of  fresh 
air  for  the  maintenance  of  life.  "Were  it  not  for  this,  we  should 
breathe  as  well  in  the  most  filthy  prison  as  among  the  most  delightful 
pastures."  *  *  *  "jj^  fact,"  he  continues,  "where  a  fire  burns 
readily  there  we  can  easily  breathe."  Note  that  there  was  no  men- 
tion that  only  a  part  of  the  air  was  taken  up  by  the  blood.  The  com- 
mon knowledge  of  the  time  was  that  air  was  a  simple  substance,  not 
a  mixture  of  several  elements  as  we  know  it  today. 

Mayow  and  His  Researches. — ^The  next  contribution  to  the  sub- 
ject of  respiration  was  that  of  John  Mayow,  bom  in  London  in  1643. 
Mayow  was  a  lawyer  by  profession  and  science  was  his  avocation. 
Many  valued  contributions  to  medical  science  were  made  by  men 
whose  lives  were  spent  in  other  callings.  Priestly  who  discovered 
oxygen  was  a  Unitarian  minister;  Schleiden,  whose  name  is  con- 
nected with  the  cell  theory,  was  a  lawyer;  Schwann  was  a  botanist; 
Metchnikoff  is  a  biologist.  Thus  many  of  the  important  discoveries 
germain  to  medicine  were  made  by  men  whose  work  was  inspired  by 
the  fascination  of  the  subject  in  hand — ^the  avocation  of  their  leisure 
moments.  Of  Mayow  it  was  said  he  took  his  degree  in  law  and 
"became  noted  for  his  practice  therein."  Mayow's  published  works 
consisted  of  four  tracts — de  sal  nitro  et  spiritu  nitro  aero;  de  respi- 
ratione ;  de  respiratione  f eotus  in  utero  et  ovo ;  de  motu  musculari  et 
spiritibus  animalibus.  He  showed  that  it  was  not  the  whole  air 
which  was  necessary  for  respiration,  but  only  a  portion,  and  that  par- 
ticular constituent  of  the  air  which  has  since  become  known  as  oxy- 
gen. In  the  language  of  the  chemists  of  his  time,  for  he  was  essen- 
tially a  chemist,  Mayow  endeavors  to  prove  "that  this  air  which  sur- 
rounds us,  and  which,  since  by  its  tenuity  it  escapes  the  sharpness  of 
our  eyes,  seems  to  those  who  think  about  it  to  be  an  empty  space, 
is  impregnated  with  a  certain  universal  salt,  of  a  nitro-saline  nature, 
that  is  to  say,  with  a  vital,  fiery,  and  in  the  highest  degree  fermen- 
tative spirit."  The  word  "salt"  was  used  by  the  seventeenth  century 
chemist  to  designate  any  substance  not  distinctly  metallic  or  liquid. 

Mayow  sums  up  the  conditions  necessary  for  combustion;  "con- 
cerning fire  it  must  be  noted  that  for  the  ignition  of  this  it  is  neces- 
sary that  igneo-aereal  (evidently  oxygen)  particles  should  either  pre- 
exist in  the  thing  to  be  burnt  or  should  be  supplied  from  the  air. 
Gunpowder  is  very  easily  burnt  by  itself  by  reason  of  the  igneo-aereal 
particles  existing  in  it.  Vegetables  are  burnt  partly  by  means  of  the 
igneo-aereal  particles  existing  in  them,  partly  by  help  of  those 
brought  to  them  from  the  air."  This  early  chemist  recognized  the 
fact  that  in  combustion  we  have  a  chemical  combination  with  the 
substance  burnt,  and  as  a  result  an  actual  increase  in  weight.  He 
experiments  with  antimony,  which  he  bums  by  focusing  the  sun's 
rays  by  means  of  a  lens;  by  weighing  the  substance  he  finds  an  in- 
crease in  weight  which  he  attributes  the  "insertion  into  it  of  igneo- 


MAYOW  45 

aereal  particles  during-  the  calcination.  As  we  shall  see  more  than  a 
century  later  Lavoisier  arrives  at  the  same  conclusion.  But  Mayow 
did  not  stop  here.  He  proceeded  to  point  out  the  identity  of  burning 
and  breathing: 

"If  a  small  animal  and  a  lighted  candle  be  shut  up  in  the  same  vessel,  the 
entrance  into  which,  of  air  from  without  is  prevented,  you  will  see  in  a  short 
time,  the  candle  go  out;  nor  will  the  animal  long  survive  its  funeral  torch. 
Indeed,  I  have  found  by  observation  that  an  animal  shut  up  in  a  flask  together 
with  a  candle  will  continue  to  breathe  for  not  much  more  than  half  the  time 
than  it  otherwise  would,  that  is  without  the  candle.  *  *  *  The  reason  why  the 
animal  can  live  some  time  after  the  candle  has  gone  out  seems  to  be  as  follows: 
The  flame  of  the  candle  needs  for  its  maintenance  a  continuous  and  at  the  same 
time  a  sufficiently  full  and  rapid  stream  of  nitro-aereal  particles.  Whence  it 
comes  about  that  if  the  succession  of  nitro-aereal  particles  be  interrupted,  even 
for  a  moment,  or  if  these  are  not  supplied  in  adequate  quantity,  the  flame  pres- 
ently sinks  and  goes  out.  Hence,  so  soon  as  the  igneo-aereal  particles  begin  to 
reach  the  flame  scantily  and  slowly,  it  is  soon  extinguished.  For  animals,  on 
the  other  hand,  a  lesser  store  of  the  aereal  food  is  sufficient,  and  one  supplied  at 
intervals,  so  that  the  animal  can  be  sustained  by  aereal  particles  remaining  after 
the  candle  has  gone  out.  Hence  it  may  be  remarked  that  the  movements  of  the 
collapsed  lungs  not  a  little  help  towards  the  sucking  of  the  aereal  particles 
which  may  remain  in  the  said  flask,  and  towards  transferring  them  into  the 
blood  of  the  breathing  animal.  Whence  it  comes  about  that  the  animal  does  not 
perish  until  just  before  the  aereal  particles  are  wholly  exhausted.  *  *  *  We  may 
infer  that  animals  and  fire  deprive  the  air  of  particles  of  the  same  kind." 

Mayow's  account  of  the  mechanics  of  respiration  would  need  lit- 
tle or  no  revision  for  a  modern  text  book  on  physiology.  He  showed 
that  the  air  entered  the  lungs  during  respiration  solely  by  atmospheric 
pressure.  He  makes  use  of  the  experiment  whereby  a  collapsed 
bladder  is  placed  into  a  bell-jar,  the  bladder  expanding  as  the  air  in 
the  jar  is  exhausted  by  means  of  an  air  pump.  He  taught  that  in  in- 
spiration the  chest  is  enlarged  by  the  descent  and  contraction  of  the 
diaphragm  and  by  the  raising  of  the  ribs.  Mayow  further  tackles 
the  raison  d'etre  of  breathing  in  which  he  shows  that  something 
necessary  to  sustain  life  passes  from  the  air  into  the  blood.  "We 
have  no  right,"  said  he,  "to  deny  the  entrance  of  air  into  the  blood 
because  on  account  of  the  bluntness  of  our  senses  we  cannot  actually 
see  the  vessels  by  which  it  makes  its  entrance." 

These  extracts  go  to  show  how  mature  the  views  of  the  seven- 
teenth century  school  of  English  physiologists,  Boyle,  Hooke,  Lower 
and  Mayow  in  particular,  were.  Mayow  by  his  nitro-aereal  or  igneo- 
aereal  substance  evidently  meant  oxygen.  Their  work  was,  however, 
allowed  to  slumber,  until  the  scientific  path  was  retraveled  by  their 
successors  nearly  a  century  later. 

Summary  Prior  to  the  Beginning  of  the  Eighteenth  Century. — 
Van  Helmont  (1648)  had  discovered  some  of  the  properties  of  car- 
bondioxide.  He  showed  that  a  gas  was  formed  from  fermentation 
or  the  combustion  of  carbon  and  from  the  action  of  vinegar  on  cer- 
tain carbonates,  and  that  this  gas  was  incapable  of  supporting  com- 
bustion. Boyle  (1670),  as  we  have  seen,  proved  that  air  was  neces- 
sary to  the  life  of  all  animals,  even  those  which  lived  under  water. 
Bernoulli,  at  a  later  date,  showed  that  the  existence  of  aquatic  ani- 
male  depended  upon  air  held  in  solution  in  water.    Hooke  exposed 


46  PATHFINDERS  OF  PHYSIOLOGY 

the  lungs  of  a  living  animal  and  maintained  the  vital  processes  by 
means  of  artificial  respiration,  showing  that  the  vital  processes  de- 
pended upon  a  continual  change  of  air  in  the  lungs.  Fracassati  drew 
attention  to  the  fact  that  the  red  color  of  the  upper  surface  of  a  clot 
was  due  to  its  exposure  to  air.  Mayow  (1674)  advanced  the  view 
that  air  contained  a  principle  capable  of  supporting  combustion,  and 
which,  absorbed  in  respiration,  changed  venous  into  arteral  blood 
and  was  the  cause  of  heat  developed  in  animal  bodies. 

Eighteenth  Century  School. — Among  the  early  eighteenth  cen- 
tury contributors  to  our  knowledge  of  respiration  was  Stephen 
Hales,  bom  1677,  who,  by  the  way,  was  not  connected  with  the  med- 
ical profession.  He  received  his  M.  A.  degree  at  Cambridge  in  1703, 
and  Bachelor  of  Divinity  in  1711.  He  was  a  clergyman  by  profession, 
a  calling  which  he  followed  until  his  death  in  1761.  He  is  chiefly 
known  as  the  inventor  of  a  "ventilator,"  by  means  of  which  fresh  air 
was  introduced  into  jails,  mines,  hospitals,  and  ships'  holds.  Four 
years  after  the  introduction  of  Hales'  invention  into  the  Savoy  prison 
only  four  prisoners  died,  whereas  the  mortality  before  its  introduc- 
tion had  been  as  high  as  one  hundred  a  year.  Devoted  as  was  Hales 
to  the  church,  he  was  even  more  devoted  to  science.  He  was  the  first 
to  determine  blood  pressure  by  actual  experiment  on  the  living  animal. 

Next  in  chronological  sequence  is  Joseph  Black,  an  eminent  chem- 
ist born  at  Bordeaux  in  1728,  where  his  father  was  engaged  in  the 
wine  trade.  Both  parents  were  of  Scotch  descent.  In  1746  Black  en- 
tered the  University  of  Glasgow,  where  he  studied  chemistry  under 
Dr.  Cullen.  He,  however,  graduated  from  the  University  of  Edin- 
burg  in  1754.  In  a  graduation  thesis  he  proved  that  the  causticity  of 
lime  and  the  alkalis  is  due  to  the  absence  of  carbonic  acid  present  in 
limestone.  He  did  not  use  the  term  carbondioxide  but  instituted  the 
term  "fixed  air."  The  former  name  was  first  used  by  Lavoisier  in 
1748.  Black's  work  was  a  distinct  contribution  to  chemistry.  In 
1756,  he  became  professor  of  anatomy  and  chemistry  at  Glasgow,  but 
shortly  become  professor  of  the  Institutes  of  Medicine.  In  the  mean- 
time he  practised  his  profession  and  found  opportunity  for  original 
investigation.  In  1766  he  was  transferred  to  a  similar  position  in 
Edinburgh.  His  lectures  were  noted  for  their  clearness  and  what  is 
perhaps  the  best  testimonial  to  any  lecturer,  his  classes  became  the 
largest  and  best  attended  in  the  university.  Though  of  delicate  con- 
stitution, by  constant  care  he  lived  to  the  fairly  ripe  age  of  seventy- 
one. 

Black  had  been  anticipated  in  his  discovery  of  "fixed  air"  by  Van 
Helmont,  whose  researches  had  been  made  a  century  earlier.  In  other 
words,  he  had  re-discovered  the  gas  later  to  be  known  as  C02.  By 
using  clear  lime  water,  he  was  able  to  show  that  "fixed  air"  was  given 
off  in  fermentation,  in  expiration  and  that  it  was  a  product  of  burn- 
ing charcoal.  The  chemical  formula  for  clear  lime  water  is  Ca  (OH)  2, 
which  in  the  presence  of  "fixed  air,"  C02,  becomes  Calcium  Carbon- 
at,  CaCO^,  which  is  precipitated  as  chalk,  and  water  (H20).  The 
result  of  the  chemical  reaction  is,  of  course,  a  reduction  in  the  caus- 
ticity of  the  original  substance. 

I  quote  the  following  extracts  from  his  treatise  on  chemistry: 

"I  had  disicovered  that  this  particular  kind  of  air,  attracted  by  alkaline  sub- 
stances, is  deadly  to  all  animals  that  breathed  it  by  the  mouth  and  nostrils  to- 


PRIESTLY  47 

gether,  but  if  the  nostrils  were  kept  shut  I  was  led  to  believe  that  it  might  be 
breathed  in  safety.  I  found  for  example  that  when  sparrows  died  in  it  in  ten  or 
eleven  seconds,  they  would  live  in  it  three  or  four  minutes  when  the  nostrils 
were  shut  by  melted  suit.  And  I  convinced  myself  that  the  change  produced  on 
wholesome  air  "by  breathing  it  consisted  chiefly,  if  not  wholly,  in  the  conversion 
of  part  of  it  into  fixed  air.  For  I  found,  that  by  blowing  through  a  pipe  into 
lime  water,  the  lime  was  precipitated,  and  the  alkali  was  rendered  mild.  ♦  *  * 
In  the  same  year  I  found  that  fixed  air  is  the  chief  part  of  the  elastic  matter, 
which  is  formed  in  liquids  in  the  vinous  fermentation.  Van  Helmont  has  indeed 
said  this.  But  it  was  at  random  that  he  said  it  was  the  same  with  the  Grotto  del 
Cane  in  Italy  (but  he  supposed  the  identity  because  both  are  deadly),  for  he  had 
examined  neither  of  them  chemically,  nor  did  he  know  that  it  was  the  air  dis- 
engaged in  the  effervescence  of  alkaline  substances  with  acids.  I  convinced 
myself  of  the  fact  by  going  to  a  brew  house  with  two  phials,  one  filled  with  dis- 
tilled water  and  the  other  with  lime  water.  I  emptied  the  first  into  a  vat  of 
wort  fermenting  briskly,  holding  the  mouth  of  the  phial  close  to  the  surface  of 
the  wort.  I  then  poured  some  of  the  lime  water  into  it,  shut  it  with  my  finger, 
and  shook  it.    The  lime  water  became  turbid  immediately." 

Black  goes  on  to  criticise  Van  Helmont's  pronouncements  as 
mere  chance  statements.  He,  himself,  verified  all  his  conclusions  by 
repeated  experiment. 

As  Black  re-discovered  under  the  term  "fixed  air"  that  which 
Van  Helmont  had  recognized  a  century  before,  so  Mayow's  igneo- 
aereal  salt  or  spirit  was  re-discovered  by  Priestly  and  Lavoisier. 

Priestly  and  His  Dephlogisticated  Air:  With  the  name  of  Joseph 
Priestly,  perhaps  more  than  any  other,  is  associated  in  the  modern 
mind  the  discovery  of  oxygen,  though  he  did  not  make  use  of  the 
term.    Of  him  Frederick  Harrison  has  said: 

"If  we  choose  one  man  as  a  type  of  the  intellectual  energy  of  the  eighteenth 
century  we  could  hardly  find  a  better  than  Joseph  Priestly,  though  his  was  not 
the  greatest  mind  of  the  century.  His  versatility,  eagerness,  activity  and  hu- 
manity; the  immense  range  of  his  curiosity  in  all  things  physical  and  social;  his 
place  in  science,  in  theology,  in'  philosophy  and  in  politics;  his  peculiar  relation 
to  the  Revolution,  and  the  pathetic  story  of  hisi  unmerited  sufferings,  may  make 
him  the  hero  of  the  eighteenth  century." 

He  was  bom  near  Leeds,  England,  in  1733,  and  died  in  the  United 
States  in  1804.  His  boyhood  was  uneventful.  His  family  was  de- 
scribed as  "simple,  sober,  honest.  God-fearing  folk,  staunch  Calvinists 
and  deeply  religious."  The  son  inherited  these  qualities  and  entered 
the  ministry  as  a  Unitarian  preacher,  an  act  which  was  particularly 
offensive  to  the  orthodoxy  of  the  time.  Benjamin  Franklin,  to  whom 
Priestly  is  endebted  for  the  incentive  for  scientific  study,  refers  to 
him  in  a  letter  as  an  "honest  heretic."  And  continuing  in  Franklin's 
charactersitic  style,  he  says: 

"I  do  not  call  him  honest  by  way  of  distinction,  for  I  think  all  the  heretics 
I  have  known  have  been  virtuous  men.  They  have  the  virtue  of  fortitude,  or 
they  would  not  venture  to  own  their  heresy;  and  they  cannot  afford  to  be  dif- 
fident in  any  of  the  other  virtues,  as  that  would  give  advantage  to  their  many 
enemies;  and  they  have  not  like  orthodox  sinners,  such  a  number  of  friends  to 
excuse  or  justify  them.  Do  not,  however,  mistake  me.  It  is  not  to  my  good 
friend's  heresy*  that  I  impute  his  honesty.  On  the  contrary  'tis  his  honesty  that 
has  brought  upon  him  the  character  of  heretic." 


48  PATHFINDERS  OF  PHYSIOLOGY 

Priestly  was  thirty  years  old  when  Franklin  was  sixty.  Priestly 
like  Franklin  was  well  informed  on  a  variety  of  subjects.  He  wrote 
learnedly  on  politics,  religion  and  on  science,  particularly  on  pneu- 
matic chemistry.  Boswell  dubbed  him  a  "literary  Jack-oi- all- trades," 
and  he  was  busy  with  proof  sheets  until  the  day  of  his  death.  His 
pamphlets  on  politics  and  religion  were  so  much  opposed  by  the  orth- 
odox theologians  of  his  day  that  they  ansv/ered  his  arguments  by 
burning  his  house  and  dispoiling  his  belongings,  a  peculiar  way  that 
the  so-called  orthodox  theology  has  had  in  the  past  of  dealing  wih 
those  bold  intrepid  spirits  who  have  dared  to  stand  for  what  they 
believed  to  be  the  truth.  His  home  surroundings  in  Birmingham 
became  so  unpleasant  that  in  self-defense  he  set  sail  for  America, 
here  to  breathe  the  atmosphere  of  civil  and  religious  freedom.  He 
was  offered  the  professorship  of  chemistry  in  the  University  of  Phila- 
delphia, but  the  following  year  moved  to  Northumberland,  a  town  on 
the  Susquehanna,  a  hundred  and  thirty  miles  northwest  of  Philadel- 
phia. He  lived  and  worked  until  his  death,  which  occurred  in  Feb- 
ruary, 1804. 

Priestly  endeavored  to  change  back  to  its  original  condition,  air 
that  had  been  breathed,  or  which  had  failed  to  support  the  flame  of  a 
candle.  He  eventually  succeeded  by  means  of  vegetation.  First  he 
experimented  by  placing  a  sprig  of  mint  into  a  glass  jar  standing  in- 
verted over  a  vessel  of  water.  Parenthetically,  Priestly  invented  the 
pneumatic  trough,  which  has  been  found  so  convenient  in  experiment- 
ing with  gases.  When  the  sprig  of  mint  had  been  growing  some 
months,  the  air  within  the  vessel  would  not  extinguish  a  flame  nor 
act  deleteriously  to  small  animals,  such  as  the  mouse,  placed  therein. 
The  growing  plant  really  contributed  to  the  flame  or  the  animal  that 
was  placed  in  the  vessel.  Further  experiment  showed  that  a  growing 
plant  placed  in  a  vessel  in  which  a  flame  had  been  extinguished  would 
in  time  render  the  atmosphere  in  the  jar  capable  of  supporting  either 
flame  or  animal  life.  This  lead  him  to  conclude:  "That  plants,  in- 
stead of  affecting  the  air  in  the  same  manner  with  animal  respiration, 
reverse  the  effects  of  breathing  and  tend  to  keep  the  atmosphere 
sweet  and  wholesome  when  it  is  become  noxious  in  consequence  of 
animals  either  living  and  breathing  or  dying  and  putrifying  in  it." 

Priestly's  researches  might  have  been  more  fruitful  in  results 
had  he  not  been  dominated  by  the  phlogiston  theory,  a  term  devised 
by  Stahl.  Phlogiston,  from  phlogistos,  burnt,  was  a  hypothetical 
principle  of  fire  regarded  as  a  material  substance.  Every  combustible 
substance  was  a  compound  of  phlogiston  and  the  phenomenon  of  com- 
bustion was  due  to  a  separation  of  the  compound  into  its  component 
elements. 

Priestly  was  able  to  obtain  the  same  gas  by  heating  mercuric 
oxide,  and  from  red  precipitate.  But  he  could  not  get  away  from 
the  phlogiston  theory.  Air  supported  combustion  because  it  took  up 
phlogiston  given  out  by  the  burning  body.  Common  air  supported 
combustion  in  proportion  as  it  was  free  from  phlogiston.  He  pre- 
pared oxygen  in  1774,  that  is  he  discovered  that  the  gas  he  prepared 
was  part  of  the  common  air,  which  supported  life  and  combustion. 
Venous  blood  was  blood  laden  with  phlogiston.  Blood  exposed  to  de- 
phlogisted  air  gave  up  its  phlogiston  and  became  bright  arterial  blood. 


LAVOISIER  49 

Some  idea  of  the  scope  of  Priestley's  researches  may  be  inferred 
from  the  mere  catalogue  of  his  discoveries.  He  is  credited  with  dis- 
covering dephlogisticated  air  (oxygen)  hydrochloric  acid,  sulphur 
dioxide,  nitrosulphuric  acid,  sulphuretted  hydrogen,  and  the  isolation 
of  amonia  gas. 

Lavoisier  and  His  Work. — Antoine  Laurent  Lavoisier  was  bom 
in  Paris  in  1742,  ten  years  later  than  the  date  on  which  Priestly  first 
saw  the  light  of  day.  As  scientist  his  career  was  practically  contem- 
poraneous with  that  of  Priestly,  who  made  the  same  momentous 
discovery,  working  independently.  In  1775,  a  year  after  Priestly  had 
prepared  his  dephlogisticated  air  (oxygen),  Lavoisier  published  his 
paper  "On  the  nature  and  principle  which  combines  with  metals  dur- 
ing their  calcination."  In  this  paper  he  showed  that  metals  on  being 
"burnt"  did  not  give  up  phlogiston  to  the  air  but  took  something  from 
the  air;  they  on  becoming  metallic  oxides,  increased  in  weight.  La- 
voisier dealt  the  death  blow  to  the  phlogiston  theory  and  was  in  a 
sense  the  real  discover  of  oxygen.  He  proved  that  the  principle  which 
combined  with  metals  when  calcined  was  the  principle  of  acidity.  He 
says:  "I  shall  therefore  designate  dephlogisticated  air,  air  eminently 
respirable,  when  in  a  state  of  combination  or  fixedness  by  the  name  of 
'acidifying  principle,'  or,  if  one  prefers  the  same  meaning  in  a  Greek 
dress,  by  that  of  'oxygine'  principle."  Lavoisier  discovered  oxygen 
and  gave  it  the  name  by  which  it  will  henceforth  be  known.  He  made 
further  experiments  in  connection  with  respiration  which  he  con- 
cluded to  be  "a  combustion,  slow  it  is  true,  but  otherwise  perfectly 
similar  to  the  combustion  of  charcoal."  He  eventually  saw,  however, 
that  some  of  the  oxygen  inspired  had  other  use  than  the  production 
of  carbon  dioxide. 

It  was  not,  however,  until  the  early  decades  of  the  nineteenth 
century  that  the  view  that  oxidation  took  place  in  the  lungs  gave 
way  to  the  accurate  theory  of  tissue  respiration.  In  1837,  Gustave 
Magnus  proved  that  both  venous  and  arterial  blood  contained  oxygen 
and  carbon  dioxid. 

Hydrogen  was  discovered  by  Cavendish  in  1781,  when  he  also 
discovered  the  composition  of  water.  Nitrogen  was  discovered  in 
1772  by  Rutherford.  Oxygen  was  prepared  by  Priestly  in  1774  and 
recognized  by  Lavoisier  the  following  year.  Carbonic  acid  gas,  or  car- 
bon dioxide  was  first  discovered  by  Van  Helmont  in  1640  and  redis- 
covered and  defined  by  Black  in  1757. 


CHAPTER  VI. 


THE  NERVOUS  SYSTEM. 

The  progress  of  knowledge  of  the  nervous  system  has  been  very 
slow.  Most  of  the  other  viscera  were  known  to  the  ancients  before 
the  brain  was  recognized.  The  word  "brain"  is  not  to  be  found  in  the 
Bible.  The  ancient  Hebrews  evidently  looked  upon  the  heart  as  the 
seat  of  the  soul.  The  kidneys  were  the  habitation  of  the  mind,  while 
the  tender  emotions  were  referred  to  the  bowels.  Plato  was  perhaps 
the  first  to  assign  the  supreme  seat  of  the  mind  to  the  brain,  but  his 
views  were  purely  speculative,  inasmuch  as  he  confounded  the  sub- 
stance of  the  brain  and  of  the  spinal  cord  with  the  marrow  of  bones. 
Aristotle,  about  335  B.  C,  examined  the  brain  for  himself  and  con- 
cluded that  its  function  had  nothing  whatever  to  do  with  the  mind, 
but  that  it  was  a  refrigerating  organ  which  cooled  the  blood  for  the 
heart.  He  reasoned  according  to  the  knowledge  of  his  time.  The 
brain  was  apparently  an  insensible  and  inexcitable  organ  as  contrast- 
ed with  the  heart,  which  is  the  opposite.  Hippocrates  recognized  how 
soon  animals  became  unconscious  from  the  loss  of  blood,  or  how 
changed  by  blood  poison  or  by  the  heated  blood  of  fever;  hence  the 
inference  by  Aristotle  that  the  conscious  mind  resided  in  the  blood 
and  that  the  great  central  organ,  the  heart,  was  the  seat  of  the  soul. 
The  arteries  (from  the  etymology,  air  tubes  or  wind  pipes)  found 
empty  after  death,  were  supposed  to  carry  air  or  "ethereal"  spirits  to 
the  rest  of  the  body.  It  was  this  great  blunder  that  delayed  for  cen- 
turies, virtually  until  Harvey's  time,  all  progress  of  knowledge  of  the 
true  function  of  the  heart.  Hippocrates  maintained  that  the  brain 
was  a  gland.  With  this  supposition  subsequent  writers  ventured  the 
suggestion  that  the  brain  secretion  was  a  subtile  fluid  which  they 
designated  "animal  spirits."  The  authority  of  such  names  as  Hip- 
pocrates and  Aristotle  forbade  first  hand  investigation  for  fully  five 
centuries.  It  must  not  be  overlooked,  however,  that  amid  all  this 
guessing,  Alcmaeon  (about  500  B.  C),  an  anatomist  and  physiologist, 
taught  that  the  brain  was  the  seat  of  the  mind  and  that  all  sensation 
traveled  to  the  brain  by  means  of  the  nerves.  He  spoke  of  the  nerves 
as  "tendons"  which  misconception  held  sway  until  Descartes,  the 
philosopher,  showed  the  difference  between  tendons  and  nerves. 

About  300  B.  C.  sprung  up  the  Alexandrian  school  of  anatomists 
and  physiologists  of  whom  Herophilus  and  Erastistratus  were  chief 
who  dissected  the  brain  and  traced  to  it  the  nerves  as  Alcmaeon  had 
done.  They  even  went  so  far  as  to  distinguish  nerves  of  sensation 
and  nerves  of  motion,  but  were  still  hampered  by  Alcmaeon's  "ten- 
dons." "When  Greece  fell  under  the  subjection  of  Alexander,  mind  went 
into  exile,  and  its  first  asylum  was  the  city  of  the  conqueror."  Under 
royal  patronage  the  study  of  anatomy  and  physiology  and  surgery 
made  great  progress.  Galen  spoke  of  Herophilus  and  Erastistratus  as 
possessing  more  accurate  knowledge  of  the  human  body  than  any  one 
before  their  time.  Herophilus  was  the  first  anatomist  of  importance 
in  the  annals  of  medicine.    He  is  said  to  have  discovered  the  lacteal 


THOMAS  WILLIS  &1 

vessels,  and  the  construction  of  the  eye,  including  the  retina.  Galen 
speaks'  of  Herophilus  as  having  a  very  intimate  knowledge  of  the 
anatomy  of  the  nervous  system.  The  term,  "torcular  Herophili" 
signifies  the  "press"  or  dilation  at  the  junction  of  the  superior  longi- 
tudinal, lateral  and  occipital  sinuses  first  described  by  Herophilus. 
Herophilus  and  his  associates  performed  vivisection  upon  condemned 
criminals.  Not  only  did  medicine  progress  during  this  early  period 
(about  300  B.  C),  but  literature,  philosophy,  mathematics,  natural 
history  and  astronomy  flourished  as  well  under  the  patronage  of 
Ptolemy.  A  great  part  of  the  record  of  this  fruitful  period  was  lost 
during  the  seventh  century  of  the  Christian  Era,  with  the  destruction 
of  the  great  Alexandrian  library. 

"The  Brain  the  seat  of  Thought  and  Sensation:"  Galen,  A.  D. 
160,  overthrew  Aristotle's  theory  in  regard  to  the  brain  and  showed 
it  to  be  the  seat  of  thought  and  sensation.  Aretaeus  (Ij^  A.  D.) 
taught  that  the  brain  controlled  the  muscular  movements  of  the  body 
by  means  of  nerves  originating  in  the  brain.  He  recognized  the 
crossing  of  the  nerves  so  that  injury  to  one  hemisphere  pro- 
duced paralysis  on  the  opposite  side.  If  injury  occurred  m  the 
cord  below  the  medulla  the  paralysis  was  on  the  same  side  as  the  in- 
jury.   The  seat  of  the  soul  was,  however,  in  the  heart. 

Andreas  Vesalius  (1514-1564)  declared  that  the  "brain  in  ap- 
propriate structures,  and  in  organs  properly  subserving  its  work 
manufactures  the  animal  spirit  which  is  by  far  the  brightest  and 
most  delicate,  and  indeed  is  a  quality  rather  than  a  natural  thing. 
*  *  *  Nerves  serve  the  same  purpose  to  the  brain  that  the  great 
artery  does  to  the  heart."  The  nerves  he  regarded  as  the  "busy  at- 
tendants and  messengers  of  the  brain."  Vesalius,  however,  is  free  m 
the  use  of  such  terms  as  "vital  soul,"  "vital  spirits,"  "animal  spirits 
which  meant  so  much  to  the  physiologist  of  his  day  and  so  little  to 
us  of  the  twentieth  century.  While  these  meaningless  terms  make  a 
great  deal  of  his  work  unintelligible,  yet  there  abound  throughout 
gleams  of  truth  as  we  understand  it  today.  He  showed  that  by  sever- 
ing a  nerve  or  by  ligation  it  was  possible  to  abolish  the  action  ot 
the  nerve  upon  the  muscle.  Regarding  the  bram  he  says :  But  how 
the  brain  performs  its  functions  in  imagination,  m  reasoning,  m 
thinking  and  in  memory,  I  can  form  no  opinion  whatever." 

Nearly  a  century  later  we  come  to  the  conclusions  of  von  Hel- 
mont  and  of  Descartes  which  were  much  less  to  the  point  than  the  ex- 
pressed opinions  of  Vesahus.  One  placed  the  seat  of  the  soul  m  the 
pylorus ;  the  other  in  the  pineal  gland. 

Malpighi  devoted  much  attention  to  the  histology  of  the  nervous 
system  but  said  practically  nothing  about  the  functions  of  the  nerves. 

Thomas  Willis:  Perhaps  the  most  important  investigator  of 
the  seventeenth  century  into  the  anatomy  and  physiology  of  the 
nervous  system  was  Thomas  Willis.  He  was  born  m  Wiltshire,  Eng- 
land in  1621,  educated  at  Oxford  where  he  graduated  with  the  degree 
of  M.  A.,  1642.  He  eventually  entered  upon  the  study  of  medicine 
and  on  graduation  was  appointed  to  a  professorial  chair  m  Oxford. 
Here  he  taught,  practised  medicine  and  pursued  his  scientific  re- 
searches.   In  1666  he  located  in  London  where  in  the  language  of  a 


52  PATHFINDERS  OP  PHYSIOLOGY 

contemporary  "he  became  so  noted  and  so  infinitely  resorted  to,  that 
never  any  physician  before  went  beyond  him  or  got  more  money  year- 
ly than  he."  Willis  possessed  a  practical  knowledge  of  the  structure 
and  functions  of  the  brain,  both  in  health  and  disease.  His  name  to- 
day is  familiar  to  all  students  of  anatomy  in  the  "circle  of  Willis," 
which  designates  the  combined  arterial  structure  at  the  base  of  the 
brain.  Sir  Michael  Foster  is  inclined  to  depreciate  the  work  of  Willis. 
The  value  of  his  book  is  much  above  the  worth  of  the  author.  It  ap- 
pears that  Willis'  thirst  for  fame  was  much  greater  than  his  love  for 
truth.  Richard  Lower,  a  contemporary  was  the  real  man  of  science 
of  his  day.  Willis  is  said  to  have  appropriated  the  work  of  Lower 
and  other  earnest  men  and  to  have  published  it  as  his  own.  Through- 
out the  work  of  this  period  we  still  have  to  deal  with  the  "corporeal 
soul,"  "animal  spirits,"  "sensitive  soul"  and  similar  phrases. 

Muscle  Irritability:  Frances  Glisson,  an  Englishman,  born,  1597, 
came  upon  the  truth  of  the  relation  of  nervous  influence  to  muscular 
contraction.  Educated  at  Cambridge,  he  became  a  Fellow  and  lec- 
turer in  Greek  in  his  Alma  Mater.  On  the  publication  of  Harvey's 
work,  in  1628,  Glisson  determined  to  turn  his  attention  to  medicine, 
and  six  years  later  he  received  his  M.  D.  degree.  He  did  not  go 
abroad  as  Harvey  did  but  pursued  his  medical  studies  in  London.  He 
was  soon  appointed  Regius  Professor  of  physic  at  Cambridge,  but  it 
seems  did  not  spend  much  time  there,  as  the  social  atmosphere  was 
not  congenial  to  him.  Cambridge  was  strongly  Royalist,  while  Glis- 
son was  a  very  pronounced  Presbyterian.  He  served  in  a  professional 
capacity  in  London  during  the  great  plague  of  1665.  He  died  at  the 
age  of  eighty  years.  He  is  probably  best  known  for  his  work  on  the 
liver.  His  name  is  familiar  to  us  in  connection  with  the  capsule  cov- 
ering that  viscus.  Glisson's  studies  on  the  liver  lead  him,  to  his  dis- 
covery regarding  the  peculiar  properties  of  muscle  tissue. 

Explaining  how  the  bile  is  discharged  into  the  intestine  only 
when  it  is  needed,  he  shows  that  the  secretion  is  greater  when  the 
gall  bladder  and  passages  are  irritated,  hence  they  must  possess  the 
power  of  being  irritated.  For  this  peculiar  property  he  suggests  the 
term  irritability.  The  idea  was  not  seized  by  contemporary  physi- 
ologists, hence  Glisson's  work  remained  dormant  until  the  following 
century,  when  Haller  made  use  of  the  term,  and  since  his  day  it  has 
become  established  in  physiology  and  has  played  an  important  part  in 
the  development  of  both  physiology  and  pathology. 

Goll  and  Phrenology:  A  name  which  has  received  but  slight  at- 
tention at  the  hands  of  biographers  is  that  of  Franz  Joseph  Gall  or 
Goll,  best  known  as  the  founder  of  the  pseudo  science  of  phrenology, 
or  "bumpology"  as  it  has  been  contemptiously  called.  Goll  was  born 
in  1758.  He  took  his  degree  in  medicine  at  the  University  of  Vienna, 
in  1785,  where  his  studies  on  brain  and  mind  began.  He  was  an  acute 
observer  of  phenomena  and  from  a  collation  of  observed  facts  was 
the  first  to  demonstrate  that  the  brain  was  the  organ  of  the  whole 
mind.  The  modem  phrenologist  with  whom  we  are  more  or  less 
familiar,  is  a  disciple  of  Goll;  his  name  will  be  remembered  as  as- 
sociated with  the  discovery  of  certain  areas  in  the  spinal  cord.  Goll 
died  in  Paris  in  1828. 


BELL  AND  MAGBNDIB  53 

Bell  and  Magendie:  One  of  the  greatest  names  in  connection 
with  the  anatomy  and  physiology  of  the  nervous  system  is  that  of 
Sir  Charles  Bell.  In  fact,  his  discovery  has  been  placed  in  importance 
in  the  same  class  as  that  of  William  Harvey.  Charles  Bell  was  born 
at  Edinburgh,  Scotland,  in  1774.  After  graduating  from  the  Uni- 
versity of  Edinburgh  he  began  the  study  of  medicine  under  his  elder 
brother  John,  who  had  already  achieved  distinction  as  anatomist. 
After  graduating  he  devoted  himself  to  anatomy  and  surgery.  He 
eventually  moved  to  London,  where  he  worked  into  a  very  lucrative 
surgical  practice.  His  first  published  work  (1798)  bore  the  cumber- 
some title  ,and  it  was  the  custom  of  writers  of  the  time  to  preface 
their  work  with  a  sentence  descriptive  of  its  contents,  of  "A  System 
of  Dissections  Explaining  the  Anatomy  of  the  Human  Body,  the 
Manner  of  Displaying  its  parts  and  their  varieties  in  Disease."  Four 
years  later  Bell  published  a  series  of  engravings  of  original  drawings 
showing  the  brain  and  nervous  system.  His  drawings  are  worthy  of 
special  mention.  His  skill  as  anatomical  artist  rivaled  that  of  anato- 
mist. He  was  also  the  author  of  a  work  entitled  **The  Anatomy  of 
Expression,"  the  object  of  which  was  to  describe  the  arrangement 
by  which  the  influence  of  the  mind  is  propagated  to  the  musculature 
of  the  face  and  to  give  a  rational  explanation  of  the  muscular  move- 
ments which  accompany  the  various  emotions  and  passions.  He 
emphasized  to  the  physician  and  surgeon  the  importance  of  a 
knowledge  of  facial  expression  in  diagnosis,  to  ascertain  the  nature 
and  extent  of  bodily  suffering.  In  these  days  of  the  clinical  labora- 
tory and  multifarous  other  clinical  methods,  the  ability  to  make  a 
diagnosis  by  observation  alone  which  amounted  to  intuition  with  the 
old-time  clinicians,  is  a  lost  art.  This  work,  which  was  illustrated 
by  himself,  had  a  wide  circulation  in  his  day. 

Charles  Bell's  most  important  work,  however,  was  the  discovery 
of  the  double  system  of  nerves  issuing  from  the  spinal  cord.  He  dis- 
covered that  in  the  nerve  trunks  are  special  sensory  filaments  to 
transmit  impressions  from  the  periphery  of  the  body  to  the  sen- 
sorium  and  motor  filaments  to  convey  motor-impressions  from  the 
brain  or  other  nerve  centres  to  muscle.  He  demonstrated  that  the 
anterior  roots  of  the  spinal  cord  were  motor  and  the  posterior  roots 
sensory. 

While  in  London,  he  was  Professor  of  Anatomy,  Physiology  and 
Surgery  in  the  College  of  Surgeons.  He  was  knighted  by  William 
IV.  He  returned  to  Edinburgh  1836  where  he  became  professor  of 
anatomy  and  surgery.  His  name  is  associated  with  the  disease 
which  he  was  the  first  to  accurately  describe,  paralysis  of  the  sev- 
enth nerve— "Bell's  Palsy."    He  died  in  1842. 

A  name  of  only  less  importance  than  that  of  Sir  Charles  Bell  is 
that  of  Magendie.  Magendie  has  been  considered  the  greatest  phy- 
sician France  had  produced  down  to  his  day.  His  work  on  physiology 
written  while  in  his  early  thirties  was  almost  immediately  translated 
into  English  and  German.  It  was  a  valuable  work,  inasmuch  as  it 
was  based  upon  experimentation.  He  was  the  first  continental  in- 
vestigator to  discover  the  function  of  the  spinal  nerves,  and  accord- 
ing to  Gorton,  contributed  more  to  the  knowledge  of  the  nervous 
system  than  any  of  his  distinguished  predecessors.    Magendie  was 


54  PATHFINDERS  OF  PHYSIOLOGY 

bom  at  Bordeaux,  France,  in  1783,  studied  medicine  in  Paris,  where 
he  became  demonstrator,  and  eventually  professor  of  anatomy  in  the 
College  of  France.    He  died  in  1855. 

Magendie  is  described  as  being  abrupt  in  manner,  even  to  rude- 
ness. His  brusque  manner  has  been  referred  to  in  his  relations  with 
his  understudy,  Claude  Bernard.  He  seems,  however,  to  have  been 
a  brilliant  if  not  very  methodical  worker.  He  refers  to  himself  as 
a  ragpicker  by  the  dust  heap  of  science.  His  work  on  the  nervous 
system  was  parallel  with  that  of  Sir  Charles  Bell,  and  the  scope  of 
the  work  of  both  is  epitomized  in  the  well-known  Bell  and  Magendie 
Law  to  the  effect  that  the  spinal  roots  may  be  divided  into  afferent 
and  efferent,  the  anterior  roots  carrying  impulses  only  from  the 
spinal  cord  to  the  periphery,  while  the  posterior  roots  carry  impulses 
from  the  periphery  to  the  central  nervous  system ;  a  nerve  fibre  can- 
not be  both  motor  and  sensory;  we  may  have  both  nerve  fibres  in  a 
single  nerve  trunk  but  the  fibres  in  each  case  are  isolated  and  con- 
duct impulses  only  in  one  or  other  direction. 

To  Claude  Bernard,  associated  with  Magendie  in  the  College 
of  France,  we  owe  the  discovery  of  the  vaso-motor  nerves. 

Broca  and  the  "Speech  Center." — In  1861  Paul  Broca,  an  immi- 
nent French  surgeon,  proved  that  there  is  a  definite  locality  in  the 
brain  which  is  the  seat  of  articulate  speech.  This  is  known  today  as 
"Broca's  Convolution."  Nine  years  later,  thanks  to  the  labors  of  such 
men  as  Hitzig,  Ferrier  and  Charcot,  it  was  shown  that  each  of  the 
special  senses  has  its  anatomical  seat  in  the  brain.  It  was  also  found 
that  each  volutary  muscle  or  group  of  muscles  could  be  made  to  con- 
tract by  the  excitation  of  certain  "centers"  or  localities  in  the  surface 
of  the  brain.  Regarding  later  progress  in  brain  physiology,  Gorton 
says:  "It  is  worth  while  to  note  the  stride  anatomy  has  made  dur- 
ing the  closing  years  of  the  nineteenth  century,  especially  in  knowl- 
edge of  the  central  nervous  system  of  man  and  animals.  Early  in  the 
last  decade  of  the  century  the  subject  was  taken  up  by  German  and 
Italian  anatomists,  Waldeyer,  Nissl,  Marchi,  Golgi,  His,  Apathy  and 
others.  To  Waldeyer  we  are  indebted  for  the  doctrine  of  neuron  as 
applied  to  nerve  cells,  from  the  Greek  word  "neuron,'  signifying  unit. 
According  to  this  doctrine  every  cell  is  a  unit  having  an  independent 
existence,  distinct  and  apart  from  other  cells,  though  related  to  them, 
and  may  degenerate  and  die  without  affecting  the  existence  of  the 
others.  Meynart  estimates  that  "the  cortex  of  the  cerebral  hemis- 
pheres alone  contains  twelve  hundred  millions  of  ganglionic  cells;" 
and  Donaldson  states  that  three  thousand  million  cells  "is  a  modest 
estimate  of  the  total  number  of  these  neurons  in  the  central  nervous 
system."  The  doctrine  of  neurons  has  been  assailed  as  applied  to  com- 
parative histology  by  the  distinguished  Apathy,  and  defended  among 
others  by  Barker,  of  Johns  Hopkins  University."  The  invention  of 
staining  processes  afforded  a  powerful  impetus  to  the  study  of  nerve 
tissues. 

By  means  of  animal  experimentation  Flourens,  Luciani  and 
Horsley  determined  the  function  of  the  cerebrum.  Removal  of  the 
cerebrum  from  a  frog  or  pigeon  caused  all  its  voluntary  movements 
to  cease,  but  did  not  interfere  with  the  reflexes  or  the  negative  func- 


PATHOLOGIC  STATES  OP  THE  BRAIN  55 

tions.  The  same  investigators  found  that  if  the  cerebrum  was  re- 
moved and  the  cerebellum  left,  the  animal  has  sense  of  appreciation, 
but  fails  in  muscular  coordination.  Stephen  Hales  showed  that  the 
spinal  cord  is  necessary  for  reflex  movements  and  Marshall  Hall  work- 
ed out  the  whole  problem  of  reflexes.  Galvani  (1791)  studied  reflexes 
by  applying  electric  stimuli  to  frogs'  legs. 

Pathologic  States  of  Brain  and  Nervous  System — Apropos  of  the 
development  of  knowledge  of  the  physiology  of  the  nervous  system  is 
the  evolution  of  our  knowledge  of  its  pathologic  states.  The  insane 
have  suffered  much  owing  to  ignorance  and  misconception  on  the 
part  of  the  sane.  Ancient  nations  looked  upon  the  insane  as  possess- 
ed of  evil  spirits  or  as  "possessed  of  devils."  Later  the  Greek,  Alex- 
andrian, and  the  Roman,  looked  upon  the  insane  man  as  a  sick  man 
and  he  was  accordingly  treated  by  means  of  drugs,  baths,  exercise  and 
other  hygienic  measures.  A  great  retrogression  took  place  during  the 
second  or  third  centuries  of  the  Christian  era.  Theories  of  demoniac 
possession  again  held  sway,  with  the  result  that  the  insane  were  sub- 
jected to  the  utmost  cruelty.  This  attitude  continued  throughout  the 
Middle  Ages.  In  fact,  no  marked  advance  was  made  until  the  eigh- 
teenth century.  Various  places  of  custody  were  maintained  for  the 
insane  where  they  were  confined  in  dungeons,  badly  clothed  and  bad- 
ly fed.  The  first  real  advances  in  their  care  were  made  by  Phihp 
Pinel,  in  France.  Tuke,  in  England,  and  Benjamin  Rush,  of  America, 
near  the  end  of  the  eighteenth  century.  Pinel  in  1793  substituted  a 
system  of  non-restraint  and  humane  treatment  for  blows  and  punish- 
ments. William  Tuke,  member  of  the  Society  of  Friends,  was  mak- 
ing similar  reforms  in  England.  Stahl,  early  in  the  eighteenth  cen- 
tury, insisted  on  the  essentially  sinful  character  of  insanity  and  this 
attitude  found  echo  in  Heinroth  in  the  early  nineteenth  century. 
Religious  theories  have  little  by  little  given  place  to  physiological  and 
psychological  explanations  until  today  the  insane  man  is  regarded  as 
a  sick  man.  Insanity  implies  disease  organic  or  functional,  just  as 
do  other  abnormal  manifestations. 

Note:    I   am   indebted  to  F.   X.  Dercum's   work    on   Mental   Diseases,    1913, 
for  the  data  of  the  last  paragraph. 


CHAPTER  VII 


THE  CELL  THEORY 

"The  cell  theory  furnishes  the  starting  point  for  all  modem 
studies  in  biology  and  enables  all  students  to  speak  the  same 
language,"  says  a  twentieth  century  writer.  The  recognition  of  the 
fact  that  animals  and  plants  are  constructed  on  a  similar  plan  must 
be  placed  among  the  most  important  discoveries  of  the  nineteenth 
century  prolific  as  that  century  has  been  in  scientific  achievement. 
"No  other  biological  generalization,"  says  Professor  Wilson, 
referring  to  the  cell  theory,  "save  only  the  theory  of  or- 
ganic evolution  has  brought  so  many  diverse  phenomena 
under  a  common  point  of  view,  or  has  accomplished  more  for  the 
unification  of  knowledge."  By  the  term  "cell-theory"  is  understood 
the  teaching  that  all  animal  and  plant  tissues  are  composed  of  units 
known  as  "cells,"  which  term  as  we  shall  see  is  inappropriate  so  far  as 
the  actual  things  designated  by  it  are  concerned.  The  cell-theory  is 
a  generalization  which  places  animals  and  plants  on  a  basis  of  similar- 
ity of  structure. 

Anticipated  in  the  Seventeenth  Century:  The  cell  doctrine  was 
anticipated  as  far  back  as  the  seventeenth  century,  for  it  is  to  a 
worker  of  the  mid-seventeenth  century  that  we  are  endebted  for  the 
term  "cell."  Robert  Hooke,  an  English  microscopist,  experimented 
with  cork,  which  he  declared  to  be  made  u-p  of  "little  boxes  or  'cells' 
distinguished  from  one  another."  He  made  thin  sections  by  means 
of  a  pen  knife  and  found  them  to  be  all  "cellular  or  porous  in  the  man- 
ner of  a  honeycomb."  Malpighi  and  Leeuwenhoek,  in  the  seventeenth 
century,  made  drawings  which  have  been  preserved  showing  the  cell 
structure  of  plants;  we  may  therefore  conclude  that  the  cell  theory 
announced  in  1838,  was  foreshadowed  by  seventeenth  century  work- 
ers. Wolff,  an  acute  scientific  observer  in  1759  worked  out  the 
identity  of  plants  and  animals,  as  shown  by  their  development.  Hux- 
ley summarizes  Wolff's  view  of  the  development  of  elementary  parts 
as  follows:  "Every  organ,  according  to  him,  is  composed  at  first  of 
a  little  mass  of  clear  viscous  nutritive  fluid  which  possesses  no  or- 
ganization of  any  kind,  but  is  at  most  composed  of  globules.  In  this 
semi-fluid  mass  cavities  are  now  developed;  these  if  they  remain 
round  or  polygonal,  become  the  subsequent  cells;  if  they  elongate, 
the  vessels ;  and  the  process  is  identically  the  same  whether  it  is  ex- 
amined in  the  vegetating  point  of  a  plant  or  the  young  budding  organs 
of  an  animal." 

Bichat's  Contribution:  Though  his  connection  with  th,e  cell 
theory  is  open  to  question,  the  name  of  Bichat  is  deserving  of  mention 
in  discussing  it.  Marie  Francois  Xavier  Bichat,  bom  in  France  in 
1771,  is  noted  as  the  founder  of  histology.  He  studied  in  Paris  under 
the  great  surgeon  Desault.  He  was  himself  made  professor  of  anatomy 
at  the  age  of  twenty-six  years,  a  position  which  he  held  until  death 
relieved  him  of  his  labors  at  the  early  age  of  thirty-one.    It  is  related 


M.  SCHLEIDEN, 


THEODOR  SCHWANN, 
Co-founders  of  the  Cell-Doctrine.     From  Locy  Biology  and  Its   Makers. 


THE  CELL  THEORY,  1838  67 

that  he  won  the  attention  and  admiration  of  his  chief  by  making  a 
complete  extemporaneous  report  of  one  of  Desault's  lectures.  Bichat 
was  a  most  admirable  character;  he  has  been  described  as  of  "mid- 
dling stature,  with  an  agreeable  face,  lighted  by  piercing  and  express 
sive  eyes,"  and  as  being  "in  all  relations  of  life  most  aimable,  a 
stranger  to  envy  or  other  hateful  passions,  modest  in  demeanour,  and 
lively  in  his  manners  which  were  open  and  free."  Two  of  his  works, 
his  treatise  on  the  membranes  and  his  general  anatomy  are  important 
as  the  foundation  of  histology,  or  the  minute  anatomy  of  the  tissues. 
After  the  ennunciation  of  the  cell  theory  Bichat's  work  took  on  a  new 
phase,  namely  that  of  microscopic  study  of  the  tissues.  Schwann's 
cell  theory  was  in  reality  an  extension  of  his  work.  Bichat's  claim 
for  credit  in  connection  with  the  cell  theory  has  been  called  into  ques- 
tion inasmuch  as  his  investigations  were  done  without  the  aid  of  the 
microscope. 

The  Cell  Theory,  1838:  During  the  first  three  decades  of  the 
Nineteenth  century  there  accumulated  a  great  mass  of  unconnected 
observations  on  the  microscopic  structure  of  both  animals  and  plants. 
"We  must  clearly  recognize,"  said  Tyson,  "the  fact  that  for  some 
time  prior  to  1838  the  cell  had  come  to  be  quite  universally  recognized 
as  a  constantly  recurring  element  in  vegetable  and  animal  tissues, 
though  little  importance  was  attached  to  it  as  an  element  of  organiza- 
tion, nor  had  its  character  been  clearly  determined." 

Eighteen  hundred  and  thirty-eight  was  an  epochal  year  in  biolog- 
ical science,  chronicling  as  it  does  the  ennunciation  of  the  cell-theory 
l9y  Schleiden  and  Schwann  the  result  of  the  combined  efforts  of 
botanist  and  animal  biologist.  The  work  of  Schwann,  however,  was 
more  comprehensive  and  important  than  that  of  Schleiden,  and  to 
him,  therefore,  belongs  the  greater  honor. 

M.  Schleiden  was  educated  for  the  legal  profession  and  had  engaged 
in  the  practice  of  law.  He  soon  abandoned  it  for  medicine,  but  after 
graduation  devoted  himself  to  the  study  of  botany.  Locy  describes 
his  work  in  1837,  stating  that  he  arrived  at  a  new  view  in  regard  to 
the  origin  of  plant  cells.  This  new  view  though  founded  upon  er- 
roneous observations  and  conclusions  served  to  provoke  discussion. 
His  work  acted  like  a  ferment,  we  are  told,  in  bringing  about  new  ac- 
tivity. Schleiden  was  noted  for  his  alertness  in  entering  upon  con- 
troversies, a  trait  which  better  befits  the  lawyer  than  the  man  of 
science  whose  sole  concern  should  be  the  quest  of  truth.  His  replies 
to  his  adversaries  were  at  times  vitriolic  and  he  often  indulged  in  bit- 
ter personalities.    Perhaps  his  legal  training  was  responsible  for  this. 

His  methods  of  investigation  were  sound,  based  as  they  were  on 
experiment  and  observation.  He  conceived  the  necessity  of  studying 
the  development  of  plants  in  order  to  understand  their  anatomy  and 
physiology.  The  nucleus  of  the  plant  cell  was  discovered  in  1831,  by 
Robert  Brown.  Schleiden  seized  upon  the  nucleus  as  the  starting 
point  of  new  cells  but  wrongly  supposed  that  the  new  cells  started 
from  a  small  clear  bubble  on  one  side  of  the  nucleus.  And  yet  it  was 
through  these  inaccurate  observations  of  Schleiden  that  his  co-found- 
er, Schwann,  arrived  at  his  general  conclusions.  An  incident  is  re- 
lated of  the  two  dining  together  one  October  evening  when  Schleiden 


58  PATHFINDERS  OF  PHYSIOLOGY 

took  occasion  to  relate  to  his  friend  his  observations  and  inferences. 
Schwann  was  impressed  at  once  with  the  similarity  to  his  own  ob- 
servations on  animal  tissues.  They  at  once  proceeded  to  Schwann's 
laboratory  where  sections  of  the  spinal  cord  were  examined.  Schleiden 
recognized  the  nuclei  as  similar  to  those  he  had  found  in  plant  cells. 

Theodor  Schwann;  Schleiden  and  Schwann  seem  to  have  been  the 
most  diverse  personalities.  The  former  was  pugnacious  and  always 
ready  to  take  up  the  gauntlet  in  controversy;  the  latter  was  one  of 
the  mildest  of  men.  We  are  endebted  to  Henle,  a  name  familiar  in 
microscopic  anatomy,  for  what  we  know  of  the  life  of  Schwann.  This 
is  Henle's  description  of  him:  "He  was  a  man  of  stature  below  the 
medium,  with  a  beardless  face,  an  almost  infantile  and  always  smiling 
expression,  smooth  dark  brown  hair,  wearing  a  fur  trimmed  dressing 
gown,  living  in  a  poorly  lighted  room  on  the  second  floor  of  a  restau- 
rant which  was  not  even  of  the  second  class.  He  would  pass  whole 
days  there  without  going  out,  with  a  few  rare  books  around  him,  and 
numerous  glass  vessels,  retorts,  vials  and  tubes,  simple  apparatus 
which  he  himself  made.  Or  I  go  in  imagination  to  the  dark  and 
fusty  halls  of  the  anatomical  institute  where  we  used  to  work  till 
night  fall  by  the  side  of  our  excellent  chief,  Johann  Muller.  We  took 
our  dinner  in  the  evening,  after  the  English  fashion  so  that  we  might 
enjoy  more  of  the  advantages  of  daylight." 

Johann  Muller:  The  mention  of  Johann  Muller  is  worth  a  mo- 
ment's digression.  Muller,  the  son  of  a  poor  shoemaker,  was  born  at 
Coblentz  in  July,  1801.  Perhaps  it  was  the  meagerness  of  his  worldly 
possessions,  for  have  not  all  the  followers  of  Saint  Crispin  been  men 
of  lowly  estate,  that  served  to  bring  out  the  true  metal  of  his  charac- 
ter. Surmounting  the  disadvantages  and  lack  of  opportunity  of 
youth  he  became  eventually  one  of  the  great  teachers  and  master 
minds  of  German  science.  The  inspiration  derived  from  a  great 
teacher  or  personality  is  difficult  to  comprehend  much  less  to  explain. 
Harvey  was  influenced  by  his  association  with  Fabricius;  Bernard 
was  similiary  inspired  by  Magendie.  The  dominant  physiological  mind 
during  the  first  half  of  the  nineteenth  century  was  that  of  Muller. 
He  was  the  great  trainer  of  anatomists  and  physiologists.  Among 
desciples  during  his  professorship  at  Berlin  were  Virchow,  the  patho- 
logist; Du  Bois  Reymond  and  Brucke,  the  physiologsits ;  Henle,  the 
anatomist;  Helmholtz,  and  Leiberkuhn.  All  became  distinguished 
scholars  and  professors  in  German  universities.  In  glowing  tribute 
to  his  master,  Helmholtz  said:  "Whoever  comes  in  contact  with  men 
of  the  first  rank  has  an  altered  scale  of  values  in  life.  Such  intellec- 
tual contact  is  the  most  interesting  event  that  life  can  offer." 

Muller's  manner  and  gestures  in  the  classroom  reminded  his 
hearers  of  a  Catholic  priest.  The  way  he  impressed  the  scientific  men 
of  his  time  is  best  evidenced  by  the  numerous  tributes  accorded  his 
memory.  Verworn  says:  "He  is  one  of  those  monumental  figures 
that  the  history  of  every  science  brings  forth  but  once.  They  change 
the  whole  aspect  of  the  field  in  which  they  work  and  all  later  growth 
is  influenced  by  their  labors."  And  of  his  monumental  work  the 
Handbook  of  Physiology,  which  appeared  in  1833,  the  same  eulogist 
writes :  "This  work  stands  today  unsurpassed  in  the  genuinely  philos- 


JOHANN  MULLER  59 

ophical  manner  in  which  the  material,  swollen  to  vast  proportions  by 
innumerable  special  researches  was  for  the  first  time  sifted  and  elab- 
orated into  a  unitary  picture  of  the  mechanism  within  the  living  or- 
ganism. In  this  respect  the  handbook  is  not  only  unsurpassed  but 
unequalled." 

To  sum  up  and  to  sift  the  accumulated  knowledge  of  a  depart- 
ment of  scientific  endeavor  is  truly  a  herculean  task,  one  requiring 
the  impartiality  of  a  judge  and  energy  and  zeal  for  the  work  that 
amounts  to  genius.  Haller  performed  a  similar  service  for  physiology 
in  his  day. 

Johann  MuUer  a  "vitalist:"  Attempts  have  been  made  to  ac- 
count in  some  more  or  less  satisfactory  way  for  the  phenomena  of  life. 
Two  theories  have  engaged  the  attention  of  scientists — vitalism  and 
the  chemico — physic  or  mechanistic  theory.  The  majority  of  scien- 
tists of  the  present  day  maintain  that  living  organisms  are  mere 
machines,  as  opposed  to  the  theory  of  vitalism  which  presupposed  the 
presence  of  some  "life"  principle.  The  chemico-physicist  today  sees 
nothing  that  may  not  be  explained  by  the  ordinary  laws  of  physics 
and  chemistry.  The  tendency  in  all  science  is  to  express  the  less  sim- 
ple in  terms  of  the  more  simple.  Every  activity  of  living  substance 
is  accompanied  by  molecular  or  chemical  changes  in  its  composition, 
such  as  oxidation  (combustion)  so  that  chemical  activity,  which  is 
the  source  of  energy,  and  all  vital  manifestations  are  physico-chemical 
in  nature.  Haller,  in  1700,  defined  vitalism  or  vital  force  as  a  life 
principle  which  possessed  the  ability  to  originate  energy,  which  meant 
that  an  organism  was  not  wholly  dependent  upon  the  food  which  it 
consumed  for  its  energy. 

The  scientists  of  the  period  1810  to  1850  were,  for  the  most  part, 
adherents  to  the  mechanical  explanation  of  the  phenomena  of  life. 
During  this  time  also,  the  vitalistic  theory  was  not  without  its  advo- 
cates who  were  among  the  pupils  of  the  great  idealist  philosopher, 
Schelling.  Such  men  as  Johann  Muller,  the  physiologist;  Von  Baer, 
the  embryologist,  and  Liebig,  the  chemist,  were  said  to  be  close  ad- 
herents to  the  vitalistic  theory.  It  was  not,  however,  until  1847,  the 
date  of  publication  of  the  researches  of  Helmholtz  on  conservation  of 
energy  that  vitalism  received  a  stunning  blow.  Sir  Michael  Foster 
explains  Muller's  vitalistic  leanings  by  declaring  that,  "He  was  a 
vitalist  only  in  the  sense  that  he  was  theoretically  of  opinion  that 
even  when  the  physico-chemical  analysis  of  vital  phenomena  had 
been  pushed  as  far  as  it  could,  there  would  still  remain  a  large  residue 
which  could  not  be  explained  by  any  such  analysis,  however  complete." 
In  view  of  the  fact  that  his  great  pupils  were  noted  for  their  effort  to 
solve  physiological  problems  by  physico-chemical  means,  the  explana- 
tion is  plausible.  It  might  be  stated  that  Schwann,  as  well  as  other 
pupils  of  Muller,  had  recourse  to  vitalistic  explanations  only 
when  their  means  of  analysis  proved  inefficient. 

"The  graven  image,  vitalism,"  says  Starling,  "has  acted  as  a  con- 
tinual check  on  the  growth  of  man's  knowledge  and  control  of  his 
environment  just  as  the  hypothesis  of  special  creation  would  impede 
all  research  into  the  relationships  of  animals  and  plants,  so  vitalism 
would  stay  the  hand  of  the  physiologist  in  his  endeavors  to  determine 
the  changes  which  occur  within  the  living  organism." 


60  PATHFINDERS  OF  PHYSIOLOGY 

Muller  died  in  1857.  Virchow,  at  his  obsequies  in  Berlin,  indulged 
in  the  following  panegyric  over  his  master  : 

"My  feeble  powers  have  been  invoked  to  honor  this  great  man, 
whom  we  all,  representatives  of  the  great  medical  family,  teachers  and 
taught,  practitioners  and  investigators,  mutually  lament  and  whose 
memory  is  still  so  vividly  with  us.  Neither  cares  by  day  nor  labors 
by  night  can  efface  from  our  mind  the  sorrow  which  we  feel  for  his 
loss.  If  the  will  made  the  deed,  how  gladly  would  I  attempt  the  hope- 
less task  of  proper  appreciation.  Few  have  been  privileged,  like  my- 
self, to  have  this  great  master  beside  them  in  every  stage  of  de- 
velopment. It  was  his  hand  which  guided  my  first  steps  as  a  medical 
student.  *  *  *  But  how  can  one  tongue  adequately  praise  a  man 
who  presided  over  the  whole  domain  of  the  science  of  natural  life ;  or 
how  can  one  tongue  depict  the  master  mind,  which  extended  the 
limits  of  his  great  kingdom  until  it  became  too  large  for  his  own  un- 
divided government  ?  *  *  *  We  have  to  inquire  what  it  was  that 
raised  Muller  to  so  high  a  place  in  the  estimation  of  his  contempor- 
aries ;  by  what  magic  it  was  that  envy  became  dumb  before  him,  and 
by  what  mysterious  means  he  contrived  to  enchain  to  himself  the 
hearts  of  beginners  and  to  keep  them  captive  through  many  long 
years  ?  Some  have  said  that  there  was  something  supernatural  about 
Muller,  that  his  whole  appearance  bore  the  stamp  of  the  uncommon. 
That  this  commanding  influence  did  not  wholly  depend  on  his  extra- 
ordinary original  endownments  is  certain  from  what  we  know  of  the 
history  of  his  mental  greatness." 

Years  of  Discovery:  Such  was  the  mind  from  which  Schwann  de- 
rived his  inspiration.  The  middle  of  the  nineteenth  century  was  the 
golden  age — ^the  Periclean  age — of  physiology  in  Germany.  To  quote 
further  from  Schwann's  biographer  (Henle) :  Those  were  great  days. 
The  microscope  had  been  brought  to  such  a  state  of  perfection  that  it 
was  available  for  accurate  scientific  observation.  The  mechanics  of  its 
manufacture  had  besides  just  been  simplified  to  such  a  degree  that 
its  cost  was  not  beyond  the  means  of  the  enthusiastic  student  even 
of  limited  means.  Any  day  a  bit  of  animal  tissue,  shaved  off  with  a 
scalpel  or  picked  to  pieces  with  a  pair  of  needles  might  lead  to  im- 
portant ground  breaking  discoveries." 

After  the  publication  of  his  work  on  the  cell  theory,  Schwann 
was  appointed  professor  in  the  University  of  Louvain,  where  he  re- 
mained nine  years,  after  which  he  received  a  similar  appointment  in 
the  University  of  Liege.  His  "Microscopical  Researches  into  the  Ac- 
cordance in  the  Structure  of  Plants  and  Animals,"  though  of  somewhat 
cumbersome  title,  is  one  of  the  great  classics  of  biology.  He  proves 
the  identity  in  structure  of  animals  and  plants  by  direct  comparison 
of  their  elementary  parts.  His  conclusion  is  that  "the  elementary 
parts  of  all  tissues  are  formed  of  cells  in  an  analogous,  though  very 
diversified  manner,  so  that  it  may  be  asserted  that  there  is  one  univer- 
sal principle  of  development  for  the  elementary  parts  of  organisms, 
however,  different  and  that  this  principle  is  the  formation  of  cells." 

Virchow  and  "Cellular"  Pathology:  Any  account  of  the  cell 
theory  must  needs  be  incomplete  with  the  omission  of  the  name  and 
work  of  Rudolph  Virchow.  Virchow  was  bom  in  1821  of  humble 
parentage,  his  father  eking  out  a  livelihood  from  the  combined  oc- 


DISCOVERY  OF  PROTOPLASM  «1 

cupations  of  farmer  and  small  shopkeeper.  The  son  who  received  the 
academic  training  of  his  day  was  of  an  active  restless  temperament. 
Virchow's  was  a  mind  open  to  new  ideas,  of  liberal  and  mdependent 
views  on  medicine,  politics  and  religion.  His  open  sympathies  with 
the  reform  tendencies  in  1848  were  such  that  he  was  obliged  to  leave 
Berlin  for  Wurzburg,  where  he  taught  pathology  and  did  much  orig- 
inal work  therein.  He  was  recalled  to  Berlin  m  1856,  when  he  was 
made  professor  of  pathology  in  the  university.  The  scope  of  his  ac- 
tivities may  be  seen  when  it  is  considered  that  he  was  also  a  member 
of  the  Reichstag,  where  he  became  leader  of  the  opposition  and  a 
vigorous  antagonist  of  Bismark.  As  chairman  of  the  finance  com- 
mittee, Virchow  is  credited  as  the  author  of  the  Prussian  Budget 
system.  He  took  a  leading  part  in  the  politics  of  his  city ;  and  the 
fact  that  from  being  one  of  the  most  unsanitary  cities  Berlin  has 
come  to  be  one  of  the  most  healthful  spots  has  been  attributed  in 
great  measure  to  his  insistance  on  sanitary  reform.  Virchow  stands 
in  much  the  same  relation  to  pathology  as  Schwann  to  histology.  He 
has  been  called  the  "Father  of  Modern  Pathology."  He  established 
"The  true  and  fertile  doctrine  that  every  morbid  structure  consists  ot 
cells  which  have  been  derived  from  pre-existing  cells,"  or  as  he  him- 
self expressed  it:  "Omnis  cellula  e  cellula  "  His  chief  work  was  his 
cellular  pathology  published  in  1858;  m  it  he  applied  the  cell  theory 
to  diseased  tissues.    He  died  in  1903. 

The  ceU  theory  incomplete  as  first  announced:  When  William 
Harvey  published  his  discovery  of  the  circulation,  so  complete  was 
Ss  lelfippointed  task  that  little  was  left  for  future  w^^jk^^^-^J^,^, 
Xcoffenic  function  of  the  liver  is  known  and  understood  by  us 
Sactfcanv  as  prodaimed  by  Claude  Bernard.  The  cell  doctrme  has  a 
fast  rd  LrenThistory.  As  announced  by  it's  cO;founders  it  was 
far  from  being  complete.  Among  other  inaccuracies  they  attached 
too  much  importance  to  the  cell  wall.  The  word  "cell"  implies  a  wall- 
ed enclosure.  The  cell  of  honeycomb  or  the  cell  of  a  penal  institution 
are  examples  which  suggest  themselves.  The  fundamental  declara- 
tion that  all  parts  of  plants  and  animals  are  built  of  similar  units  or 
structures  has  been  substantiated.  This  is  perhaps  the  only  portion 
of  the  theory  that  has  not  been  profoundly  changed. 

The  Discovery  of  Protoplasm:  Perhaps  of  equal  importance  to 
the  cell-theory  was  the  recognition  of  protoplasm.  Huxley  called  it 
"the  physical  basic  of  life."  Fehx  Dujardin  recognized  this  sub- 
stance which  is  the  basis  of  vital  activity,  m  1835.  He  discovered  m 
lower  animal  forms  a  jelly-like  substance  which  he  called  sarcode 
Dujardin  was  born  in  1801  at  Tours,  France.  He  was  trained  to  fol- 
low the  trade  of  his  father,  namely,  that  of  watchmaker,  and  the 
manual  dexterity  thus  acquired  served  him  in  good  stead  m  the  later 
vocation  of  his  life.  He  was  an  adept  with  the  microscope  and  pos- 
sessed no  small  ability  as  sketch  artist.  He  showed  early  a  love  for 
the  natural  sciences.  His  contributions  to  science  cover  a  range  of 
topics.  He  was  perhaps  the  greatest  authority  of  his  day  on  proto- 
zoology.   He  died  in  1860. 

Schleiden  saw  protoplasm  but  called  it  gum.  Cohn,  in  1850, 
taught  that  "protoplasm"  of  plants  and  "sarcode"  of  lower  aniinal 
life  were  the  same  thing.    Max  Schultze,  in  1861,  confirmed  Cohn  s 


62  PATHFINDERS  OP  PHYSIOLOGY 

position  and  added  that  the  cell  consisted  of  little  units  of  protoplasm 
surrounding  a  nucleus.  The  nucleus  was  first  described  by  Fontana, 
in  1871.  It  was  regarded  as  a  normal  element  of  the  cell  by  Robert 
Brown  in  1883.  It  was  eventually  seen  that  many  cells,  especially 
animal  cells,  are  without  a  ceil  wall,  hence  the  conclusion  that  the  so- 
called  "wall"  is  not  an  essential  feature  of  the  "cell."  When  the  cell 
wall  is  absent  the  protoplasm  is  the  cell.  The  nucleus  was  found  to 
be  within  the  substance  of  the  cell  and  not  within  the  cell  wall. 
Schultze  defined  the  cell  as  a  globule  of  protoplasm  surrounding  a 
nucleus.  From  being  regarded  as  an  element  of  structure  merely,  the 
cell  has  come  to  be  recognized  as  the  physiological  unit  within  which 
all  physiological  activity  takes  place. 

Perhaps  the  most  authoritative  as  well  as  the  most  recent  defi- 
nition of  protoplasm  is  the  following  significant  paragraph  by  Star- 
ling: 

"Though  it  may  be  convenient  to  have  a  word  such  as  protoplasm  signifying 
simply  'living  material,'  it  is  important  to  remember  there  is  no  such  thing  as  a 
single  substance — protoplasm.  The  reactions  of  every  cell  as  well  as  its  organiza- 
tion are  the  resultant  of  the  molecular  structure  of  matter  of  which  it  is  built  up. 
The  gross  methods  of  the  chemist  show  him  that  the  composition  of  the  proto- 
plasm of  the  muscle  cell  is  entirely  different  from  that  of  a  leucocyte  or  white 
blood  corpuscle.  The  finer  methods  of  the  physiologist  show  him  that  every  sort 
of  cell  in  the  body  has  its  own  manner  of  life,  its  own  peculiarities  of  reaction 
to  uniform  changes  in  its  surroundings.  No  individual  will  react  in  exactly  the 
same  manner  as  another  individual  even  of  the  same  species,  and  the  reactions 
of  the  whole  organism  are  but  the  sum  of  the  reactions  of  it's  constituent  cells. 
There  is  not  one  protoplasm  therefore,  but  an  infinity  of  protoplasms  and  the  use 
of  the  term  can  be  justified  only  if  we  keep  this  fact  in  mind  and  use  the  word 
merely  as  a  convenient  abbreviation  for  any  material  endowed  with  life.  Even 
in  a  single  cell  there  is  more  than  one  kind  of  protoplasm.  In  its  chemical 
characters,  in  its  mode  of  life,  and  in  its  reactions,  the  nucleus  differs  widely 
from  the  cytoplasm.  Both  are  necessary  to  the  life  of  the  cell  and  both  must 
be  regarded  according  to  our  present  ideas  as  living.'  In  the  cytoplasm  itself 
we  find  structures  or  substances  which  we  must  regard  as  on  their  way  to  proto- 
plasm or  as  products  of  the  break  down  of  protoplasm;  but  in  many  cases  it  is 
impossible  to  say  whether  a  given  material  is  to  be  regarded  as  lifeless  or  as  re- 
active living  matter.  Even  in  a  single  cell  we  may  have  differentiation  among 
its  different  parts,  one  part  serving  for  the  process  of  digestion  while  other  parts 
are  employed  for  purpose  of  locomotion.  Here  again  there  must  be  chemical 
differences,  and  therefore  diflerent  protoplasms." 

A  statement  of  the  cell  theory  at  the  present  time  (1913)  must 
include  four  conceptions:  (1)  The  cell  as  a  unit  of  structure;  (2)  The 
cell  as  a  unit  of  physiological  activity;  (3)  The  cell  as  embracing  all 
hereditary  qualities  within  its  substance;  (4)  The  cell  in  the  histori- 
cal development  of  the  organism." 

Students  of  cytology  have  sought  to  find  out  if  any  uniformity 
of  organization  of  protoplasm  exists.  Accordingly  we  have  a  number 
of  explanations  or  theories  regarding  its  structure.  Altmann  pro- 
posed the  granular  theory.  By  the  employment  of  certain  hardening 
reagents  he  demonstrated  dense  masses  of  spherical  or  rod-shaped 
granules  in  all  the  cells  of  the  body.  In  these  he  located  the  various 
vital  functions,  the  sum  total  of  which  constitute  the  life  of  the  cell. 


THE  NUCLEUS  63 

He  further  maintained  that  these  granules  could  come  only  by  division 
of  pre-existing  granules.  He  parodied  Virchow's  famous  phrase 
omnis  cellula  e  cellula  into  omne  granulum  e  granulo. 

The  fibrillar  theory  presupposes  net-work  or  clusters  of  fibrils 
known  as  "spongio-plasm"  (sponge  plasm)  in  contra -destinction  to 
clear  or  structureless  matter  filling  in  the  meshes  of  the  net  to  which 
the  name  "hyaloplasm  (glass  plasm)  has  been  given. 

In  the  Alveolar  Theory  of  Butschli  the  author  regards  the  so- 
called  granules  as  products  manufactured  by  the  hyaline  protoplasm 
and  stored  up  as  spherules  so  that  the  protoplasm  between  the  drop- 
lets form  an  alveolar  partition — hence  the  name  of  the  theory. 

Discussing  the  question  as  to  the  fluidity  of  protoplasm  Starling 
regards  it  as  "essentially  fluid  in  character,  the  form  and  rigidity 
which  are  acquired  by  most  cells  being  due  to  chemical  and  physical 
differentiation  occurring  in  its  fluids." 

The  cell  consists  of  cytoplasm  and  nucleus.  Cytoplasm  (cell  plasm) 
is  a  term  formulated  by  Kolliker  in  1863.  Though  not  so  applied 
when  first  used,  it  has  come  to  mean  the  living  substance  of  the  cell 
body  other  than  the  nucleus.  Cytoplasm  contains,  for  the  most  part, 
substances  apparently  foreign  to  the  cell  proper.  In  the  cytoplasm 
of  plant  cells,  for  example,  are  stored  up  starches  and  oils.  Most 
nerve  cells  contain  various  shaped  bodies  which,  it  is  alleged,  repre- 
sent stored  up  energy.  The  passive  bodies  in  the  cytoplasm  are  sup- 
posed to  represent  some  form  of  latent  energy  upon  which  the  cell 
may  draw.  In  the  cells  of  any  green  leaf  are  to  be  found  spherical 
masses  which  play  a  most  important  role  in  the  lives  of  not  only  plants 
but  of  animals  as  well.  By  the  action  of  the  sun's  rays  a  chemical 
change  takes  place  in  these  bodies  known  to  botanists  as  chloroplasts 
by  which  carbondioxide  and  water  are  broken  down,  decomposed  and 
immediately  synthetized  into  a  different  substance — carbohydrate, 
starch,  which  will  respond  to  the  well  known  iodine  test  for  starch. 
Carbohydrate  is  one  of  the  food  principles.  Fats  are  also  made  and 
stored  in  the  form  of  oils.  In  spite  of  the  fact  that  the  atmospheric 
air  surrounding  the  plant  contains  an  abundance  of  free  nitrogen. 
The  plant  cells  are  unable  to  make  use  of  it.  Nitrogen  must  be  first 
combined  as  a  nitrate,  become  dissolved  in  the  soil  and  taken  up  by 
the  roots  of  the  plants,  or  in  the  case  of  water  plants,  by  special  cells, 
before  the  green  matter  in  the  leaf  can  be  transformed  into  protein. 
The  plant,  therefore,  has  power  to  make  foods  out  of  the 
chemical  elements  of  air  and  water  when  these  elements  are 
properly  combined.  This  is  the  only  source  of  food  of  both  plant 
and  animal  and  it  is  the  result  of  cellular  activity. 

The  Nucleus:  The  nucleus  has  been  recognized  as  a  most  es- 
sential part  of  the  cell.  It  not  only  takes  part  in  the  complex  process 
of  cell  division  but  dominates  the  rest  of  the  cell.  It  is  not  my  pur- 
pose to  enter  upon  a  discussion  of  the  morphology  and  physiology  of 
the  animal  and  vegetable  cell,  further  than  it  is  necessary  to  trace  the 
various  stages  of  the  history  of  its  revelation  from  its  earliest  recog- 
nition to  the  present.  The  reader  is  referred  to  the  numerous  excel- 
lent text  books  on  the  subject. 


64 


PATHFINDERS  OF  PHYSIOLOGY 


ILLUSTRATIONS  SHOW  DIAGRAMATICALLY  THE  CELL  AND 
INDIRECT  CELL  DIVISION. 


Fig.  1. 


Fig.  2. 


Fig.  3. 


Fig.  4. 


Fig.  5. 


Fig.  6. 


Fig.  7. 


.  'y^^,  first  change  in  the  appearance  of  the  nucleus  which  indicates  that  a 
division  is  about  to  take  place,  consists  in  a  rearrangement  of  the  chromatin  net 
work,  which  now  takes  place  on  the  appearance  of  a  tangled  thread  (Fig.  2). 
The  outwardly  directed  loops  of  this  skein  often  correspond  to  the  seperate  por- 
tions into  which  the  thread  eventually  breaks  up.  The  thread  gradually  grows 
shorter  and  thicker,  and  presently  becomes  divided  into  a  number  of  pieces  known 
as  chromosomes.  In  the  chromosomes  the  shortening  and  thickening  process  is 
continued  until  these  bodies  arrive  finally  at  the  form  of  stumpy  rods,  each  of 
which,  often  becomes  bent  into  the  form  of  a  horse  shoe.  Meanwhile  the  nuclear 
membrane,  breaks  down,  so  that  the  hyaline  substance  of  the  nucleus  becomes 
continuous  with  that  of  the  cell  body  surrounding  it.  A  fresh  phenomenon  now 
becomes  visible.  A  spindle-shaped  arrangement  makes  it's  appearance  consisting 
of  a  number  of  minute  fibrils  which  connect  together  two  points — the  poles  of 
the  spindle — situated  at  opposite  ends  of  the  cell.  The  chromosomes  now  change 
their  position  so  that  they  come  to  be  in  the  plane  of  the  equator  of  the  spindle, 
and  about  this  line  each  chromosome  splits  longitudinally  into  two  great  por- 
tions (Fig.  4  and  5).  This  splitting  in  the  case  of  each  chromosome  takes  place 
in  the  equatorial  plane  of  the  spindle,  so  that  one  member  of  each  pair  of  daugh- 
ter chromosomes  faces  towards  one  pole  of  the  spindle  and  the  second  towards 
the  other  pole.  The  members  of  each  pair  of  daughter  chromosomes  now  begin 
to  move  away  from  one  towards  the  two  poles  of  the  spindle,  and  as  they  do  so 
the  first  indication  of  a  dividing  wall  between  the  second  new  cells  begins  to 
make  its  appearance  in  the  equatorial  plane.  Arriving  at  the  poles,  the  daughter 
chromosomes  begin  to  elongate  and  to  put  out  processes  which  finally  meet  and 
fuse  with  those  of  their  neighbors  to  form  the  chromatin  reticulum  of  the  new 
nuclei.  (Fig.  7.)  Surrounding  each  new  nucleus,  thus  developing  at  either  pole 
of  the  now  rapidly  disappearing  spindle,  a  new  nuclear  membrane  makes  it's  ap- 
pearance; the  dividing  wall  in  the  position  of  the  equator  of  the  spindle  develops 
into  a  complete  partition  in  tfie  case  of  plants.  (The  animal  cell  is  without  a 
cell  wall.)  The  division  into  two  new  cells  is  thus  completed.  (Fig.  8.)  Each 
new  cell  is  provided  with  a  nucleus  into  which  has  entered  precisely  its  fair 
share  of  the  chromatin  which  was  present  in  the  parent  nucleus." 
— Illustration  and  description  after  Locke. 


THE  CELL  IN  HEREDITY  65 

The  discovery  of  the  various  dyes  and  tissue  stains  afforded  a 
wonderful  stimulous  to  the  microscopic  study  of  tissues  as  well  as  to 
bacteriological  studies.  It  is  hard  to  conceive  of  much  progress  in 
bacteriology  without  this  aid.  Dyes  for  staining  protoplasm  were  first 
prepared  in  1868.  The  property  of  taking  up  a  stain  gave  rise  to  the 
invention  of  a  number  of  new  names  for  which  scientists  have  as 
usual  drawn  freely  from  the  Greek.  To  designate  that  protoplasm 
which  stained  deeply,  we  have  the  term  "chromatin."  The  word 
"achromatin"  has  been  applied  to  protoplasm,  which  will  not  absorb 
the  dye.  Certain  rod-shaped  bodies  situated  within  the  nucleus, 
which  stain  more  deeply  than  any  other  portions  are  known  as 
"chromosomes." 

The  Cell  In  Heredity — Within  recent  years  the  subject  of  here- 
dity has  claimed  the  attention  of  biologists  and  its  practical  applica- 
tion has  become  of  intense  interest  to  the  laity,  advances  in  our 
knowledge  of  heredity  are  already  producing  results.  They  have 
revolutionized  agricultural  methods  as  shown  in  the  marked  improve- 
ment of  animals  and  plants.  It  is  impossible  of  realization  what  are 
the  potentialities  in  regard  to  the  improvement  of  the  human  race. 
Eugenics  is  as  yet  in  its  infancy.  The  past  ten  years  has  witnessed 
the  production  of  voluminous  literature  on  eugenics  and  its  kindred 
subject  heredity. 

Smallwood  in  his  latest  work-  states  that  "Whatever  may  be 
the  ultimate  analysis  of  the  problem  of  heredity,  there  can  be  no 
hesitation  in  stating  that  the  transmitted  characters  exist  potentially 
in  the  protoplasm  of  the  cell.  From  the  egg  of  a  robin  only  a  robin 
will  develop,  from  the  ovum  of  an  oak  only  an  oak  will  grow  and  dur- 
ing the  growth  each  follows  its  own  successive  developmental  stages 
even  to  the  minutest  details.  It  has  been  well  said  'nature  never  yet 
made  two  eggs  or  two  sperms  exactly  alike.'  The  cells  which  give 
rise  to  new  organisms  are  the  germ  cells,  sperms  and  ova.  These 
differ  greatly  in  shape  and  size — some  of  the  sperm  cells  being  but 
one  one-hundred-thousandths  the  bulk  of  the  ovum  and  yet  the  pa- 
ternal characters  are  easily  recognized  in  the  adult.  *  *  *  The 
cells  of  the  body  are  divided  into  body  plasm  and  germ  plasm."  Germ 
plasm  might  be  looked  upon  as  the  immortal  in  man  in  as  much  as  it 
is  continuous.  After  the  germ  plasm  has  given  rise  to  a  new  individ- 
ual, some  of  it  is  left  behind  to  participate  in  the  formation  of  a  new 
offspring,  so  as  Davenport  puts  it,  "There  is  really  no  inheritance 
from  parent  to  child  but  parent  and  child  resemble  each  other  because 
they  are  derived  from  the  same  plasm,  they  are  chips  of  the  same 
old  block;  and  the  son  is  half-brother  of  the  father  by  another 
mother." 

As  the  cell  has  been  called  "The  physiological  unit,"  and  proto- 
plasm "the  physical  basis  of  life,"  the  chromosomes  have  been  proven 
the  physical  basis  of  heredity.  They  are  very  definite  and  import- 
ant organs.  The  number  which  make  their  appearance  at  each  cell 
division  is  the  same  in  all  the  cells  of  any  given  creature  and  is  con- 
stant for  the  cells  of  the  members  of  any  given  species. 

"The  remarkable  fact,"  says  Wilson,  "has  been  established  that 
every  species  of  plant  or  animal  has  a  fixed  and  characteristic  num- 


66  PATHFINDERS  OF  PHYSIOLOGY 

ber  of  chromosomes,  which  regularly  occurs  in  the  division  of  all  of  its 
cells,  and  in  all  forms  arising  by  sexual  reproduction,  the  number 
is  even." 

Whatever  the  offspring  is,  it  is  potential  in  the  fertilized  ovum. 
If  this  is  the  contribution  of  each  parent,  the  role  performed  by  the 
mother  is  that  of  custodian  of  her  embryonic  charge  until  birth.  Her 
power  to  alter  it  in  any  way  is  as  futile  as  that  of  the  father.  The 
parent  is  rather  the  trustee  of  the  germ  plasm  than  the  producer  of 
the  child.  Sir  Michael  Foster  once  said,  'The  animal  body  is  in  reality 
a  vehicle  for  the  ova;  and  after  the  life  of  the  parent  has  become 
potentially  renewed  in  the  offspring,  the  body  remains  as  a  cast-off 
envelope  whose  future  is  but  to  die."  The  germ  plasm  is  "the  lighted 
torch  handed  on  from  one  runner  to  another."  Et  quasi  cursores 
vital  lampada  tradunt.  This  equally  true  of  plant  life,  where  the 
plant  matures  and  dies  leaving  the  future  offspring  potentially  in  the 
seed.  How  characteristics  are  transmitted  from  ancestor  to  offspring 
is  not  known. 

NOTE: — It  has  been  estimated  that  the  number  of  cells  entering  into  the 
composition  of  the  body  of  an  adult  human  being  is  about  twenty-six  million 
five  hundred  thousand  millions   (26,500,000,000,000). 


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